{"id":382,"date":"2023-06-15T01:42:28","date_gmt":"2023-06-15T05:42:28","guid":{"rendered":"https:\/\/opentextbooks.concordia.ca\/explorations\/chapter\/12\/"},"modified":"2025-07-09T17:12:00","modified_gmt":"2025-07-09T21:12:00","slug":"12","status":"publish","type":"chapter","link":"https:\/\/opentextbooks.concordia.ca\/explorations\/chapter\/12\/","title":{"raw":"Modern Homo sapiens","rendered":"Modern Homo sapiens"},"content":{"raw":"<div class=\"__UNKNOWN__\">\r\n\r\nKeith Chan, Ph.D., Grossmont-Cuyamaca Community College District and MiraCosta College\r\n\r\n<em>This chapter is a revision from \"<\/em><a class=\"rId7\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-14\/\"><em>Chapter 12: Modern Homo sapiens<\/em><\/a><em>\u201d by Keith Chan. In <\/em><a class=\"rId8\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, first edition<\/em><\/a><em>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under <\/em><a class=\"rId9\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\"><em>CC BY-NC 4.0<\/em><\/a><em>. <\/em>\r\n<div class=\"textbox textbox--learning-objectives\"><header class=\"textbox__header\">\r\n<h2 class=\"textbox__title\"><span style=\"color: #ffffff\">Learning Objectives<\/span><\/h2>\r\n<\/header>\r\n<div class=\"textbox__content\">\r\n<ul>\r\n \t<li>Identify the skeletal and behavioral traits that represent modern <em>Homo sapiens.<\/em><\/li>\r\n \t<li>Critically evaluate different types of evidence for the origin of our species in Africa and our expansion around the world.<\/li>\r\n \t<li>Understand how the human lifestyle changed when people transitioned from foraging to agriculture.<\/li>\r\n \t<li>Hypothesize how human evolutionary trends may continue into the future.<\/li>\r\n<\/ul>\r\n<\/div>\r\n<\/div>\r\n<p class=\"import-Normal\">The walls of a pink limestone cave in the hillside of Jebel Irhoud jutted out of the otherwise barren landscape of the Moroccan desert (Figure 12.1). Miners had excavated the cave in the 1960s, revealing some fossils. In 2007, a re-excavation of the site became a momentous occasion for science. A fossil cranium unearthed by a team of researchers was barely visible to the untrained eye. Just the fossil\u2019s robust brows were peering out of the rock. This research team from the Max Planck Institute for Evolutionary Anthropology was the latest to explore the ancient human presence in this part of North Africa after a find by miners in 1960. Excavating near the first discovery, the researchers wanted to learn more about how <em>Homo sapiens<\/em> lived far from East Africa, where we thought our species originated.<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"alignnone\" width=\"2500\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/06\/image10-1.jpg\" alt=\"Rocky hillside with exposed layers. People are visible at the base.\" width=\"2500\" height=\"987\" \/> Figure 12.1: The excavation of an exposed cave at Jebel Irhoud, Morocco, where hominin fossils were found in the 1960s and in 2007. Dating showed that they could represent the earliest-known modern Homo sapiens. Credit: <a href=\"https:\/\/www.eva.mpg.de\/homo-sapiens\/presskit.html\">View looking south of the Jebel Irhoud (Morocco) site<\/a> by Shannon McPherron, <a href=\"https:\/\/www.eva.mpg.de\/index.html\">MPI EVA Leipzig<\/a>, is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/\">CC BY-SA 2.0 License<\/a>.[\/caption]\r\n\r\nThe scientists were surprised when they analyzed the cranium, named Irhoud 10, and other fossils. Statistical comparisons with other human crania concluded that the Irhoud face shapes were typical of recent modern humans while the braincases matched ancient modern humans. Based on the findings of other scientists, the team expected these modern <em>Homo sapiens<\/em> fossils to be around 200,000 years old. Instead, dating revealed that the cranium had been buried for around 315,000 years.\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Together, the modern-looking facial dimensions and the older date reshaped the interpretation of our species: modern <em>Homo sapiens<\/em>. Some key evolutionary changes from the archaic <em>Homo sapiens<\/em> (described in Chapter 11) to our species today happened 100,000 years earlier than we had thought and across the vast African continent rather than concentrated in its eastern region.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">This revelation in the study of modern <em>Homo sapiens<\/em> is just one of the latest in this continually advancing area of biological anthropology. Researchers today are still discovering amazing fossils and ingenious ways to collect data and test hypotheses about our past. Through the collective work of many scientists, we are building an overall theory of modern human origins. <span style=\"background-color: #ff99cc\">In this chapter, we will first cover the skeletal changes from archaic <em>Homo sapiens<\/em> to modern <em>Homo sapiens<\/em>. Next, we will track how modern <em>Homo sapiens<\/em> expanded around the world. Lastly, we will cover the development of agriculture and how it changed human culture.<\/span><\/p>\r\n\r\n<h2 class=\"import-Normal\">Defining Modernity<\/h2>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">What defines modern <em>Homo sapiens<\/em> when compared to archaic <em>Homo sapiens<\/em>? Modern humans, like you and me, have a set of derived traits that are not seen in archaic humans or any other hominin. As with other transitions in hominin evolution, such as increasing brain size and bipedal ability, modern traits do not appear fully formed or all at once. In other words, the first modern <em>Homo sapiens<\/em> was not just born one day from archaic parents. The traits common to modern <em>Homo sapiens<\/em> appeared in a <strong>[pb_glossary id=\"2708\"]mosaic[\/pb_glossary]<\/strong> manner: gradually and out of sync with one another. There are two areas to consider when tracking the complex evolution of modern human traits. One is the physical change in the skeleton. The other is behavior inferred from the size and shape of the cranium and material culture evidence.<\/p>\r\n\r\n<h3 class=\"import-Normal\"><strong>Skeletal Traits<\/strong><\/h3>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The skeleton of modern <em>Homo sapiens<\/em> is less robust than that of archaic <em>Homo sapiens<\/em>. In other words, the modern skeleton is <strong>[pb_glossary id=\"1406\"]gracile[\/pb_glossary]<\/strong>, meaning that the structures are thinner and smoother. Differences related to gracility in the cranium are seen in the braincase, the face, and the mandible. There are also broad differences in the rest of the skeleton.<\/p>\r\n\r\n<h4 class=\"import-Normal\"><em>Cranial Traits<\/em><\/h4>\r\n[caption id=\"\" align=\"alignleft\" width=\"445\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image29-2.png\" alt=\"A rounded skull facing a robust skull with sloping forehead.\" width=\"445\" height=\"221\" \/> Figure 12.2: Comparison between modern (left) and archaic (right) Homo sapiens skulls. Note the overall gracility of the modern skull, as well as the globular braincase. Credit: <a class=\"rId15\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-14\/\">Modern human and Neanderthal<\/a> original to <a class=\"rId16\" href=\"https:\/\/explorations.americananthro.org\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a class=\"rId17\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.[\/caption]\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Several elements of the braincase differ between modern and archaic <em>Homo sapiens<\/em>. Overall, the shape is much rounder, or more [pb_glossary id=\"1758\"]<strong>globular<\/strong>,[\/pb_glossary] on a modern skull (Lieberman, McBratney, and Krovitz 2002; Neubauer, Hublin, and Gunz 2018; Pearson 2008; Figure 12.2). You can feel the globularity of your own modern human skull. Feel the height of your forehead with the palm of your hand. Viewed from the side, the tall vertical forehead of a modern <em>Homo sapiens<\/em> stands out when compared to the sloping archaic version. This is because the frontal lobe of the modern human brain is larger than the one in archaic humans, and the skull has to accommodate the expansion. The vertical forehead reduces a trait that is common to all other hominins: the brow ridge or <strong>[pb_glossary id=\"1759\"]supraorbital torus[\/pb_glossary]<\/strong>. The parietal lobes of the brain and the matching parietal bones on either side of the skull both bulge outward more in modern humans. At the back of the skull, the archaic occipital bun is no longer present. Instead, the occipital region of the modern human cranium has a derived tall and smooth curve, again reflecting the globular brain inside.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The trend of shrinking face size across hominins reaches its extreme with our species as well. The facial bones of a modern <em>Homo sapiens<\/em> are extremely gracile compared to all other hominins (Lieberman, McBratney, and Krovitz 2002). Continuing a trend in hominin evolution, technological innovations kept reducing the importance of teeth in reproductive success (Lucas 2007). As natural selection favored smaller and smaller teeth, the surrounding bone holding these teeth also shrank.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Related to smaller teeth, the mandible is also gracile in modern humans when compared to archaic humans and other hominins. Interestingly, our mandibles have pulled back so far from the prognathism of earlier hominins that we gained an extra structure at the most anterior point, called the <strong>[pb_glossary id=\"2709\"]mental eminence[\/pb_glossary]<\/strong>. You know this structure as the chin. At the skeletal level, it resembles an upside-down \u201cT\u201d at the centerline of the mandible (Pearson 2008). Looking back at archaic humans, you will see that they all lack a chin. Instead, their mandibles curve straight back without a forward point. What is the chin for and how did it develop? Flora Gr\u00f6ning and colleagues (2011) found evidence of the chin\u2019s importance by simulating physical forces on computer models of different mandible shapes. Their results showed that the chin acts as structural support to withstand strain on the otherwise gracile mandible.<\/p>\r\n\r\n<h4 class=\"import-Normal\"><em>Postcranial Gracility<\/em><\/h4>\r\n[caption id=\"\" align=\"alignright\" width=\"368\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image16-5.png\" alt=\"Two complete skeletons. The left is taller with a thinner frame.\" width=\"368\" height=\"575\" \/> Figure 12.3: Anterior views of modern (left) and archaic (right) Homo sapiens skeletons. The modern human has an overall gracile appearance at this scale as well. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-14\/\">Modern and archaic Homo sapiens skeletons (Figure 12.3)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.[\/caption]\r\n\r\nThe rest of the modern human skeleton is also more gracile than its archaic counterpart. The differences are clear when comparing a modern <em>Homo sapiens<\/em> with a cold-adapted Neanderthal (Sawyer and Maley 2005), but the trends are still present when comparing modern and archaic humans within Africa (Pearson 2000). Overall, a modern <em>Homo sapiens<\/em> postcranial skeleton has thinner cortical bone, smoother features, and more slender shapes when compared to archaic <em>Homo sapiens<\/em> (Figure 12.3). Comparing whole skeletons, modern humans have longer limb proportions relative to the length and width of the torso, giving us lankier outlines.\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Why is our skeleton so gracile compared to those of other hominins? Natural selection can drive the gracilization of skeletons in several ways (Lieberman 2015). <span style=\"background-color: #ffff00\">A slender frame is adapted for the efficient long-distance running ability that started with <em>Homo erectus<\/em>. Furthermore, slenderness is a genetic adaptation for cooling an active body in hotter climates, which aligns with the ample evidence that Africa was the home continent of our species.<\/span><\/p>\r\n\r\n<h3 class=\"import-Normal\"><strong>Behavioral Modernity<\/strong><\/h3>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Aside from physical differences in the skeleton, researchers have also uncovered evidence of behavioral changes associated with increased cultural complexity from archaic to modern humans. How did cultural complexity develop? Two investigations into this question are archaeology and the analysis of reconstructed brains.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Archaeology tells us much about the behavioral complexity of past humans by interpreting the significance of material culture. In terms of advanced culture, items created with an artistic flair, or as decoration, speak of abstract thought processes (Figure 12.4). The demonstration of difficult artistic techniques and technological complexity hints at social learning and cooperation as well. According to paleoanthropologist John Shea (2011), one way to track the complexity of past behavior through artifacts is by measuring the variety of tools found together. The more types of tools constructed with different techniques and for different purposes, the more modern the behavior. Researchers are still working on an archaeological way to measure cultural complexity that is useful across time and place.<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"alignleft\" width=\"221\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image15-1-1.jpg\" alt=\"A brown standing statue of a human figure with cat\u2019s head.\" width=\"221\" height=\"392\" \/> Figure 12.4: Carved ivory figure called \u201cthe Lion-Man of the Hohlenstein-Stadel.\u201d It dates to the Aurignacian culture, between 35 and 40 kya. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Loewenmensch1.jpg\">Loewenmensch1<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Dagmar_Hollmann\">Dagmar Hollmann<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.[\/caption]\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The interpretation of brain anatomy is another promising approach to studying the evolution of human behavior. When looking at investigations on this topic in modern <em>Homo sapiens<\/em> brains, researchers found a weak association between brain size and test-measured intelligence (Pietschnig et al. 2015). Additionally, they found no association between intelligence and biological sex. These findings mean that there are more significant factors that affect tested intelligence than just brain size. Since the sheer size of the brain is not useful for weighing intelligence within a species, paleoanthropologists are instead investigating the differences in certain brain structures. The differences in organization between modern <em>Homo sapiens<\/em> brains and archaic <em>Homo sapiens<\/em> brains may reflect different cognitive priorities that account for modern human culture. As with the archaeological approach, new discoveries will refine what we know about the human brain and apply that knowledge to studying the distant past.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Taken together, the cognitive abilities in modern humans may have translated into an adept use of tools to enhance survival. Researchers Patrick Roberts and Brian A. Stewart (2018) call this concept the <strong>[pb_glossary id=\"1802\"]generalist-specialist niche[\/pb_glossary]<\/strong>: our species is an expert at living in a wide array of environments, with populations culturally specializing in their own particular surroundings. The next section tracks how far around the world these skeletal and behavioral traits have taken us.<\/p>\r\n\r\n<h2 class=\"import-Normal\">First Africa, Then the World<\/h2>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">What enabled modern <em>Homo sapiens<\/em> to expand its range further in 300,000 years than <em>Homo erectus<\/em> did in 1.5 million years? <span style=\"background-color: #ffff00\">The key is the set of derived biological traits from the last section. The gracile frame and neurological anatomy allowed modern humans to survive and even flourish in the vastly different environments they encountered.<\/span> Based on multiple types of evidence, the source of all of these modern humans was Africa. Instead of originating from just one location, evidence shows that modern Homo sapiens evolution occurred in a complex gene flow network across Africa, a concept called <strong>[pb_glossary id=\"1760\"]African multiregionalism[\/pb_glossary]<\/strong> (Scerri et al. 2018).<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">This section traces the origin of modern <em>Homo sapiens<\/em> and the massive expansion of our species across all of the continents (except Antarctica) by 12,000 years ago. While modern <em>Homo sapiens<\/em> first shared geography with archaic humans, modern humans eventually spread into lands where no human had gone before. Figure 12.5 shows the broad routes that our species took expanding around the world. I encourage you to make your own timeline with the dates in this part to see the overall trends.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><img class=\"aligncenter\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image13-6.png\" alt=\"315 to 195 KYA. Northern to eastern coasts of Africa are shaded.\" width=\"554\" height=\"428\" \/><\/p>\r\n<p class=\"import-Normal\"><img class=\"aligncenter\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image17-5.png\" alt=\"195-100 KYA. Africa, southern Europe and Asia are shaded\" width=\"554\" height=\"428\" \/><\/p>\r\n<p class=\"import-Normal\"><img class=\"aligncenter\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image27-3.png\" alt=\"99 to 30 KYA. Africa, Indonesia, Australia, and southern portions of Europe and Asia are shaded.\" width=\"554\" height=\"428\" \/><\/p>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"554\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image30-2.png\" alt=\"29 to 9 KYA. Shading covers most land except Antarctica, Greenland, and some islands.\" width=\"554\" height=\"428\" \/> Figure 12.5a-d: Four maps depicting the estimated range of modern Homo sapiens through time. The shaded area is based on geographical connections across known sites. Note the growth in the area starting in Africa and the oftentimes-coastal routes that populations followed. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-14\/\">Four maps depicting the estimated range of modern Homo sapiens through time<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Elyssa Ebding at <a href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\">GeoPlace, California State University, Chico<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.[\/caption]\r\n<h3 class=\"import-Normal\"><strong>Modern <\/strong><strong><em>Homo sapiens<\/em><\/strong><strong> Biology and Culture in Africa<\/strong><\/h3>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">We start with the ample fossil evidence supporting the theory that modern humans originated in Africa during the Middle Pleistocene, having evolved from African archaic <em>Homo sapiens<\/em>. The earliest dated fossils considered to be modern actually have a mosaic of archaic and modern traits, showing the complex changes from one type to the other. Experts have various names for these transitional fossils, such as <strong>[pb_glossary id=\"1763\"]<strong>Early Modern <\/strong><strong><em>Homo sapiens\u00a0 <\/em><\/strong>[\/pb_glossary][pb_glossary id=\"1764\"] or Early Anatomically Modern Humans[\/pb_glossary]<\/strong>. However they are labeled, the presence of some modern traits means that they illustrate the origin of the modern type. Three particularly informative sites with fossils of the earliest modern <em>Homo sapiens<\/em> are Jebel Irhoud, Omo, and Herto.<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"alignleft\" width=\"281\"]<img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image9-1-1.jpg\" alt=\"3D image of a human cranium with pronounced brow ridges.\" width=\"281\" height=\"282\" \/> Figure 12.6: Composite rendering of the Jebel Irhoud hominin based on micro-CT scans of multiple fossils from the site. The facial structure is within the modern human range, while the braincase is between the archaic and modern shapes. Credit: <a href=\"https:\/\/www.eva.mpg.de\/homo-sapiens\/presskit.html\">A composite reconstruction of the earliest known Homo sapiens fossils from Jebel Irhoud (Morocco) based on micro computed tomographic scans<\/a> by Philipp Gunz, <a href=\"https:\/\/www.eva.mpg.de\/index.html\">MPI EVA Leipzig<\/a>, is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/\">CC BY-SA 2.0 License<\/a>.[\/caption]\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Recall from the start of the chapter that the most recent finds at Jebel Irhoud are now the oldest dated fossils that exhibit some facial traits of modern <em>Homo sapiens<\/em>. Besides Irhoud 10, the cranium that was dated to 315,000 years ago (Hublin et al. 2017; Richter et al. 2017), there were other fossils found in the same deposit that we now know are from the same time period. In total there are at least five individuals, representing life stages from childhood to adulthood. These fossils form an image of high variation in skeletal traits. For example, the skull named Irhoud 1 has a primitive brow ridge, while Irhoud 2 and Irhoud 10 do not (Figure 12.6). The braincases are lower than what is seen in the modern humans of today but higher than in archaic <em>Homo sapiens<\/em>. The teeth also have a mix of archaic and modern traits that defy clear categorization into either group.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Research separated by nearly four decades uncovered fossils and artifacts from the Kibish Formation in the Lower Omo Valley in Ethiopia. These Omo Kibish hominins were represented by braincases and fragmented postcranial bones of three individuals found kilometers apart, dating back to around 233,000 years ago (Day 1969; McDougall, Brown, and Fleagle 2005; Vidal et al. 2022). One interesting finding was the variation in braincase size between the two more-complete specimens: while the individual named Omo I had a more globular dome, Omo II had an archaic-style long and low cranium.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Also in Ethiopia, a team led by Tim White (2003) excavated numerous fossils at Herto. There were fossilized crania of two adults and a child, along with fragments of more individuals. The dates ranged between 160,000 and 154,000 years ago. The skeletal traits and stone-tool assemblage were both intermediate between the archaic and modern types. Features reminiscent of modern humans included a tall braincase and thinner zygomatic (cheek) bones than those of archaic humans (Figure 12.7). Still, some archaic traits persisted in the Herto fossils, such as the supraorbital tori. Statistical analysis by other research teams concluded that at least some cranial measurements fit just within the modern human range (McCarthy and Lucas 2014), favoring categorization with our own species.<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"alignright\" width=\"373\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image4-3.jpg\" alt=\"Replica cranium showing wide brow ridges and gracile face.\" width=\"373\" height=\"373\" \/> Figure 12.7: This model of the Herto cranium showing its mosaic of archaic and modern traits. Credit: <a href=\"https:\/\/boneclones.com\/product\/homo-sapiens-idaltu-bou-vp-16-1-herto-skull-BH-045\/category\/all-fossil-hominids\/fossil-hominids\">Homo sapiens idaltu BOU-VP-16\/1 Herto Cranium<\/a> by <a href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a> is used by permission and available here under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.[\/caption]\r\n<p class=\"import-Normal\">The timeline of material culture suggests a long period of relying on similar tools before a noticeable diversification of artifacts types. Researchers label the time of stable technology shared with archaic types the [pb_glossary id=\"1766\"]<strong>Middle Stone Age<\/strong>[\/pb_glossary], while the subsequent time of diversification in material culture is called the[pb_glossary id=\"1767\"] <strong>Later Stone Age<\/strong>[\/pb_glossary].<\/p>\r\n<p class=\"import-Normal\">In the Middle Stone Age, the sites of Jebel Irhoud, Omo, and Herto all bore tools of the same flaked style as archaic assemblages, even though they were separated by almost 150,000 years. The consistency in technology may be evidence that behavioral modernity was not so developed. No clear signs of art dating back this far have been found either. Other hypotheses not related to behavioral modernity could explain these observations. The tool set may have been suitable for thriving in Africa without further innovation. Maybe works of art from that time were made with media that deteriorated or perhaps such art was removed by later humans.<\/p>\r\n<p class=\"import-Normal\">Evidence of what <em>Homo sapiens<\/em> did in Africa from the end of the Middle Stone Age to the Later Stone Age is concentrated in South African cave sites that reveal the complexity of human behavior at the time. For example, Blombos Cave, located along the present shore of the Cape of Africa facing the Indian Ocean, is notable for having a wide variety of artifacts. The material culture shows that toolmaking and artistry were more complex than previously thought for the Middle Stone Age. In a layer dated to 100,000 years ago, researchers found two intact ochre-processing kits made of abalone shells and grinding stones (Henshilwood et al. 2011). Marine snail shell beads from 75,000 years ago were also excavated (Figure 12.8; d\u2019Errico et al. 2005). Together, the evidence shows that the Middle Stone Age occupation at Blombos Cave incorporated resources from a variety of local environments into their culture, from caves (ochre), open land (animal bones and fat), and the sea (abalone and snail shells). This complexity shows a deep knowledge of the region\u2019s resources and their use\u2014not just for survival but also for symbolic purposes.<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"563\"]<img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image5-2-1.jpg\" alt=\"Multiple views of shells with holes bored through them.\" width=\"563\" height=\"482\" \/> Figure 12.8: Examples of the perforated shell beads found in Blombos Cave, South Africa: (a) view of carved hole seen from the inside; (b) arrows indicate worn surfaces due to repetitive contact with other objects, such as with other beads or a connecting string; (c) traces of ochre; and (d) four shell beads showing a consistent pattern of perforation. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:BBC-shell-beads.jpg\">BBC-shell-beads<\/a> by Chenshilwood (Chris Henshilbood and Francesco d\u2019Errico) at <a href=\"https:\/\/en.wikipedia.org\/wiki\/\">English Wikipedia<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.[\/caption]\r\n<p class=\"import-Normal\">On the eastern coast of South Africa, Border Cave shows new African cultural developments at the start of the Later Stone Age. Paola Villa and colleagues (2012) identified several changes in technology around 43,000 years ago. Stone-tool production transitioned from a slower process to one that was faster and made many <strong>[pb_glossary id=\"1777\"]microliths[\/pb_glossary]<\/strong>, small and precise stone tools. Changes in decorations were also found across the Later Stone Age transition. Beads were made from a new resource: fragments of ostrich eggs shaped into circular forms resembling present-day breakfast cereal O\u2019s (d\u2019Errico et al. 2012). These beads show a higher level of altering one\u2019s own surroundings and a move from the natural to the abstract in terms of design.<\/p>\r\n\r\n<h4 class=\"import-Normal\"><span style=\"background-color: #ff99cc\"><em>Summary of Modern <\/em>H. sapiens<em> in Africa<\/em><\/span><\/h4>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">The combined fossil evidence paints a picture of diversity in geography and traits. Instead of evolving in just East Africa, the Jebel Irhoud find revealed that early modern <em>Homo sapiens<\/em> had a wide range across Middle Pleistocene Africa. Supporting this explanation, fossils have different mosaics of archaic and modern traits in different places and even within the same area. The high level of diversity from just these fossils shows that the modern traits took separate paths toward the set we have today. The connections were convoluted, involving fluctuating gene flow among small groups of regional nomadic foragers across a large continent over a long time.<\/span><\/p>\r\n<p class=\"import-Normal\"><span style=\"background-color: #ff99cc\">African culture experienced a long constant phase called the Middle Stone Age until a faster burst of change produced innovation and new styles. The change was not one moment but rather an escalation in development. Later Stone Age culture introduced elements seen across many regions, including the construction of composite tools and even the use of strung decorations such as beads. These developments appear in the Later Stone Age of other regions, such as Europe. Based on the early date of the African artifacts, Later Stone Age culture may have originated in Africa and passed from person to person and region to region, with people adapting the general technique to their local resources and viewing the meaning in their own way.<\/span><\/p>\r\n\r\n<h3 class=\"import-Normal\"><strong>Expansion into the Middle East and Asia<\/strong><\/h3>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">While modern <em>Homo sapiens<\/em> lived across Africa, some members eventually left the continent. These pioneers could have used two connections to the Middle East or West Asia. From North Africa, they could have crossed the Sinai Peninsula and moved north to the [pb_glossary id=\"1769\"]<strong>Levant<\/strong>[\/pb_glossary], or eastern Mediterranean. Finds in that region show an early modern human presence. Other finds support the <strong>[pb_glossary id=\"1770\"]Southern Dispersal model[\/pb_glossary]<\/strong>, with a crossing from East Africa to the southern Arabian Peninsula through the Straits of Bab-el-Mandeb. It is tempting to think of one momentous event in which people stepped off Africa and into the Middle East, never to look back. In reality, there were likely multiple waves of movement producing gene flow back and forth across these regions as the overall range pushed east. The expanding modern human population could have thrived by using resources along the southern coast of the Arabian Peninsula to South Asia, with side routes moving north along rivers. The maximum range of the species then grew across Asia.<\/p>\r\n\r\n<h4 class=\"import-Normal\"><em>Modern <\/em>Homo sapiens<em> in the Middle East<\/em><\/h4>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Geographically, the Middle East is the ideal place for the African modern <em>Homo sapiens<\/em> population to inhabit upon expanding out of their home continent. In the Eastern Mediterranean coast of the Levant, there is a wealth of skeletal and material culture linked to modern <em>Homo sapiens<\/em>. Recent discoveries from Saudi Arabia further add to our view of human life just beyond Africa.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The Caves of Mount Carmel in present-day Israel have preserved skeletal remains and artifacts of modern <em>Homo sapiens<\/em>, the first-known group living outside Africa. The skeletal presence at Misliya Cave is represented by just part of the left upper jaw of one individual, but it is notable for being dated to a very early time, between 194,000 and 177,000 years ago (Hershkovitz et al. 2018). Later, from 120,000 to 90,000 years ago, fossils of multiple individuals across life stages were found in the caves of Es-Skhul and Qafzeh (Shea and Bar-Yosef 2005). The skeletons had many modern <em>Homo sapiens<\/em> traits, such as globular crania and more gracile postcranial bones when compared to Neanderthals. Still, there were some archaic traits. For example, the adult male Skhul V also possessed what researchers Daniel Lieberman, Osbjorn Pearson, and Kenneth Mowbray (2000) called marked or clear occipital bunning. Also, compared to later modern humans, the Mount Carmel people were more robust. Skhul V had a particularly impressive brow ridge that was short in height but sharply jutted forward above the eyes (Figure 12.9). The high level of preservation is due to the intentional burial of some of these people. Besides skeletal material, there are signs of artistic or symbolic behavior. For example, the adult male Skhul V had a boar\u2019s jaw on his chest. Similarly, Qafzeh 11, a juvenile with healed cranial trauma, had an impressive deer antler rack placed over his torso (Figure 12.10; Coqueugniot et al. 2014). Perforated seashells colored with [pb_glossary id=\"1806\"]<strong>ochre<\/strong>[\/pb_glossary], mineral-based pigment, were also found in Qafzeh (Bar-Yosef Mayer, Vandermeersch, and Bar-Yosef 2009).<\/p>\r\n&nbsp;\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"484\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image6-2-1.jpg\" alt=\"Side view of a skull replica with a globular braincase.\" width=\"484\" height=\"484\" \/> Figure 12.9: This Skhul V cranium model shows the sharp browridges. The contour of a marked occipital bun is barely visible from this angle. Credit: <a href=\"https:\/\/boneclones.com\/product\/homo-sapiens-skull-skhul-5-BH-032\">Homo sapiens Skull Skhul 5<\/a> by <a href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a> is used by permission and available here under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.[\/caption]\r\n\r\n&nbsp;\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"484\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image26-1-1.jpg\" alt=\"Human skeleton in a stony matrix. Ribs are visible below the antlers.\" width=\"484\" height=\"312\" \/> Figure 12.10 This cast of the Qafzeh 11 burial shows the antler\u2019s placement over the upper torso. The forearm bones appear to overlap the antler. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Moulage_de_la_s%C3%A9pulture_de_l'individu_%22Qafzeh_11%22_(avec_ramure_de_cervid%C3%A9),_homme_de_N%C3%A9andertal.jpg\">Moulage de la s\u00e9pulture de l'individu \"Qafzeh 11\" (avec ramure de cervid\u00e9), homme de N\u00e9andertal<\/a> (Collections du Mus\u00e9um national d'histoire naturelle de Paris, France) by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Eunostos\">Eunostos<\/a> has been modified (cropped and color modified) and is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.[\/caption]\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">One remaining question is, what happened to the modern humans of the Levant after 90,000 years ago? Another site attributed to our species did not appear in the region until 47,000 years ago. Competition with Neanderthals may have accounted for the disappearance of modern human occupation since the Neanderthal presence in the Levant lasted longer than the dates of the early modern <em>Homo sapiens<\/em>. John Shea and Ofer Bar-Yosef (2005) hypothesized that the Mount Carmel modern humans were an initial expansion from Africa that failed. Perhaps they could not succeed due to competition with the Neanderthals who had been there longer and had both cultural and biological adaptations to that environment.<\/p>\r\n\r\n<h4 class=\"import-Normal\"><em>Modern <\/em>Homo sapiens<em> of China<\/em><\/h4>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">A long history of paleoanthropology in China has found ample evidence of modern human presence. Four notable sites are the caves at Fuyan, Liujiang, Tianyuan, and Zhoukoudian. In the distant past, these caves would have been at least seasonal shelters that unintentionally preserved evidence of human presence for modern researchers to discover.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">At Fuyan Cave in Southern China, paleoanthropologists found 47 adult teeth associated with cave formations dated to between 120,000 and 80,000 years ago (Liu et al. 2015). It is currently the oldest-known modern human site in China, though other researchers question the validity of the date range (Michel et al. 2016). The teeth have the small size and gracile features of modern <em>Homo sapiens<\/em> dentition.<\/p>\r\n<p class=\"import-Normal\">The fossil Liujiang (or Liukiang) hominin (67,000 years ago) has derived traits that classified it as a modern <em>Homo sapiens<\/em>, though primitive archaic traits were also present. In the skull, which was found nearly complete, the Liujiang hominin had a taller forehead than archaic <em>Homo sapiens<\/em> but also had an enlarged occipital region (Figure 12.11; Brown 1999; Wu et al. 2008). Other parts of the skeleton also had a mix of modern and archaic traits: for example, the femur fragments suggested a slender length but with thick bone walls (Woo 1959).<\/p>\r\n&nbsp;\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"486\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image19-1-2.jpg\" alt=\"A human skull with very slight brow ridges and an extremely globular braincase.\" width=\"486\" height=\"323\" \/> Figure 12.11: The Liujiang cranium shows the tall forehead and overall gracile appearance typical of modern Homo sapiens. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Liujiang_cave_skull-a._Homo_Sapiens_68,000_Years_Old.jpg\">Liujiang cave skull-a. Homo Sapiens 68,000 Years Old<\/a> (Taken at the David H. Koch Hall of Human Origins, <a href=\"https:\/\/naturalhistory.si.edu\/visit\">Smithsonian Natural History Museum<\/a>) by <a href=\"https:\/\/www.flickr.com\/people\/14405058@N08\">Ryan Somma<\/a> has been modified (color modified) and is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/legalcode\">CC BY-SA 2.0 License<\/a>.[\/caption]\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Another Chinese site to describe here is the one that has been studied the longest. In the Zhoukoudian Cave system (Figure 12.12), where <em>Homo erectus<\/em> and archaic <em>Homo sapiens<\/em> have also been found, there were three crania of modern <em>Homo sapiens<\/em>. These crania, which date to between 34,000 and 10,000 years ago, were all more globular than those of archaic humans but still lower and longer than those of later modern humans (Brown 1999; Harvati 2009). When compared to one another, the crania showed significant differences from one another. Comparison of cranial measurements to other populations past and present found no connection with modern East Asians, again showing that human variation was very different from what we see today.<\/p>\r\n&nbsp;\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"610\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image12-1.jpg\" alt=\"A cave opening amongst a dry wooded region.\" width=\"610\" height=\"458\" \/> Figure 12.12: The entrance to the Upper Cave of the Zhoukoudian complex, where crania of three ancient modern humans were found. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Zhoukoudian_Upper_Cave.jpg\">Zhoukoudian Upper Cave<\/a> by Mutt is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.[\/caption]\r\n<h4 class=\"import-Normal\"><span style=\"background-color: #ff99cc\"><em>Summary of Modern <\/em>H. sapiens<em> in the Middle East and Asia<\/em><\/span><\/h4>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">As in Africa, the finds of the Middle East have shown that humans were biologically diverse and had complex relationships with their environment. Work in the Levant showed an initial expansion north from the Sinai Peninsula that did not last. Away from the Levant, expansion continued. Local resources were used to make lithics and decorative items.<\/span><\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">The early Asian presence of modern <em>Homo sapiens<\/em> was complex and varied as befitting the massive continent. What the evidence shows is that people adapted to a wide array of environments that were far removed from Africa. From the Levant to China, humans with modern anatomy used caves that preserved signs of their presence. Faunal and floral remains found in these shelters speak to the flexibility of the human omnivorous diet as local wildlife and foliage became nourishment. Decorative items, often found as burial goods in planned graves, show a flourishing cultural life.<\/span><\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">Eventually, modern humans at the southeastern fringe of the geographical range of the species found their way southeast until some became the first humans in Australia.<\/span><\/p>\r\n\r\n<h3 class=\"import-Normal\"><strong>Crossing to Australia<\/strong><\/h3>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Expansion of the first modern human Asians, still following the coast, eventually entered an area that researchers call <strong>[pb_glossary id=\"1772\"]Sunda[\/pb_glossary]<\/strong> before continuing on to modern Australia. Sunda was a landmass made up of the modern-day Malay Peninsula, Sumatra, Java, and Borneo. Lowered sea levels connected these places with land bridges, making them easier to traverse. Proceeding past Sunda meant navigating <strong>[pb_glossary id=\"1773\"]Wallacea[\/pb_glossary]<\/strong>, the archipelago that includes the Indonesian islands east of Borneo. In the distant past, there were many <strong>[pb_glossary id=\"864\"]megafauna[\/pb_glossary]<\/strong>, large animals that migrating humans would have used for food and materials (such as utilizing animals\u2019 hides and bones). Further southeast was another landmass called <strong>[pb_glossary id=\"1774\"]Sahul[\/pb_glossary]<\/strong>, which included New Guinea and Australia as one contiguous continent. Based on fossil evidence, this land had never seen hominins or any other primates before modern <em>Homo sapiens<\/em> arrived. Sites along this path offer clues about how our species handled the new environment to live successfully as foragers.<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"alignright\" width=\"380\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image18-1-1.jpg\" alt=\"A cranium showing a diagonal sloping forehead.\" width=\"380\" height=\"252\" \/> Figure 12.13: Replica of the Kow Swamp 1 cranium. The shape of the braincase could be due to artificial cranial modification. A competing hypothesis is that it reflects the primitive shape of Homo erectus. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Kow_Swamp1-Homo_sapiens.jpg\">Kow Swamp1-Homo sapiens<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/14405058@N08\">Ryan Somma<\/a> from Occoquan, USA, under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/legalcode\">CC BY-SA 2.0 License<\/a> has been modified (background cleaned and color modified) and is available here under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.[\/caption]\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The skeletal remains at Lake Mungo, land traditionally owned by Mutthi Mutthi, Ngiampaa, and Paakantji peoples, are the oldest known in the continent. The now-dry lake was one of a series located along the southern coast of Australia in New South Wales, far from where the first people entered from the north (Barbetti and Allen 1972; Bowler et al. 1970). Two individuals dating to around 40,000 years ago show signs of artistic and symbolic behavior, including intentional burial. The bones of Lake Mungo 1 (LM1), an adult female, were crushed repeatedly, colored with red ochre, and cremated (Bowler et al. 1970). Lake Mungo 3 (LM3), a tall, older male with a gracile cranium but robust postcranial bones, had his fingers interlocked over his pelvic region (Brown 2000).<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Kow Swamp, within traditional Yorta Yorta land also in southern Australia, contained human crania that looked distinctly different from the ones at Lake Mungo (Durband 2014; Thorne and Macumber 1972). The crania, dated between 9,000 and 20,000 years ago, had extremely robust brow ridges and thick bone walls, but these were paired with globular features on the braincase (Figure 12.13).<\/p>\r\n<p class=\"import-Normal\">While no fossil humans have been found at the Madjedbebe rock shelter in the North Territory of Australia, more than 10,000 artifacts found there show both behavioral modernity and variability (Clarkson et al. 2017). They include a diverse array of stone tools and different shades of ochre for rock art, including mica-based reflective pigment (similar to glitter). These impressive artifacts are as far back as 56,000 years old, providing the date for the earliest-known presence of humans in Australia.<\/p>\r\n\r\n<h4 class=\"import-Normal\"><span style=\"background-color: #ff99cc\"><em>Summary of Modern <\/em>H. sapiens<em> in Australia<\/em><\/span><\/h4>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">The overall view of the first modern humans in Australia from a biological perspective shows a high amount of skeletal diversity. This is similar to the trends seen earlier in Africa, the Middle East, and East Asia. The earliest-known arrivals brought with them a multifaceted suite of cultural practices as seen in their material culture.<\/span><\/p>\r\n\r\n<h3 class=\"import-Normal\"><strong>From the Levant to Europe<\/strong><\/h3>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The first modern human expansion into Europe occurred after other members of our species settled East Asia and Australia. As the evidence from the Levant suggests, modern human movement to Europe may have been hampered by the presence of Neanderthals. Another obstacle was that the colder climate was incompatible with the biology of African modern <em>Homo sapiens<\/em>, <span style=\"background-color: #ffff00\">which was adapted for exposure to high temperature and ultraviolet radiation.<\/span> Still, by 40,000 years ago, modern <em>Homo sapiens<\/em> had a detectable presence. This time was also the start of the Later Stone Age or <strong>[pb_glossary id=\"1775\"]Upper Paleolithic[\/pb_glossary]<\/strong>, when there was an expansion in cultural complexity. There is a wealth of evidence from this region due to a Western bias in research, the proximity of these findings to Western scientific institutions, and the desire of Western scientists to explore their own past. <span style=\"background-color: #ff99cc\">This section will cover key evidence of early modern human life in Europe, and the typologies used to view cultural changes in this region.<\/span><\/p>\r\n\r\n[caption id=\"\" align=\"alignleft\" width=\"323\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image11-3.jpg\" alt=\"Robust cranium with a gradually sloping forehead.\" width=\"323\" height=\"323\" \/> Figure 12.14: This side view of the Oase 2 cranium shows the reduced brow ridges but also occipital bunning that is a sign that modern Homo sapiens interbred with Neanderthals. Credit: <a href=\"https:\/\/humanorigins.si.edu\/evidence\/human-fossils\/fossils\/oase-2\">Oase 2<\/a> by James Di Loreto &amp; Donald H. Hurlbert, <a href=\"https:\/\/www.si.edu\/\">Smithsonian<\/a> [exhibit: Human Evolution Evidence, Human Fossils] has been modified (sharpened) and <a href=\"https:\/\/www.si.edu\/termsofuse\">is used for educational and non-commercial purposes as outlined by the Smithsonian.<\/a>[\/caption]\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In Romania, the site of Pe\u0219tera cu Oase (Cave of Bones) had the oldest-known remains of modern <em>Homo sapiens<\/em> in Europe, dated to around 40,000 years ago (Trinkaus et al. 2003a). Among the bones and teeth of many animals were the fragmented cranium of one person and the mandible of another (the two bones did not fit each other). Both bones have modern human traits similar to the fossils from the Middle East, but they also had Neanderthal traits. Oase 1, the mandible, had a mental eminence but also extremely large molars (Trinkaus et al. 2003b). This mandible has yielded DNA that surprisingly is equally similar to DNA from present-day Europeans and Asians (Fu et al. 2015). This means that Oase 1 was not the direct ancestor of modern Europeans. The Oase 2 cranium has the derived traits of reduced brow ridges along with archaic wide zygomatic cheekbones and an occipital bun (Figure 12.14; Rougier et al. 2007).<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Dating to around 26,000 years ago, P\u0159edmost\u00ed near P\u0159erov in the Czech Republic was a site where people buried over 30 individuals along with many artifacts. Eighteen individuals were found in one mass burial area, a few covered by the scapulae of woolly mammoths (Germonpr\u00e9, L\u00e1zni\u010dkov\u00e1-Galetov\u00e1, and Sablin 2012). The P\u0159edmost\u00ed crania were more globular than those of archaic humans but tended to be longer and lower than in later modern humans (Figure 12.15; Velem\u00ednsk\u00e1 et al. 2008). The height of the face was in line with modern residents of Central Europe. There was also skeletal evidence of dog domestication, such as the presence of dog skulls with shorter snouts than in wild wolves (Germonpr\u00e9, L\u00e1zni\u010dkov\u00e1-Galetov\u00e1, and Sablin et al. 2012). In total, P\u0159edmost\u00ed could have been a settlement dependent on mammoths for subsistence and the artificial selection of early domesticated dogs.<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"423\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image25-3.png\" alt=\"Black-and-white photograph of a human skull with labeled cranial landmarks.\" width=\"423\" height=\"389\" \/> Figure 12.15: This illustration is based upon one of the surviving photographic negatives since the original fossil was lost in World War II. The modern human chin is prominent, as is an archaic occipital bun. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:P%C5%99edmost%C3%AD_9.png\">P\u0159edmost\u00ed 9<\/a> by J. Matiegka (1862\u20131941) has been modified (sharpened) and is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.[\/caption]\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The sequence of modern <em>Homo sapiens<\/em> technological change in the Later Stone Age has been thoroughly dated and labeled by researchers working in Europe. Among them, the Gravettian tradition of 33,000 years to 21,000 years ago is associated with most of the known curvy female figurines, often assumed to be \u201cVenus\u201d figures. Hunting technology also advanced in this time with the first known boomerang, [pb_glossary id=\"1792\"]<strong>atlatl<\/strong>[\/pb_glossary] (spear thrower), and archery. The Magdalenian tradition spread from 17,000 to 12,000 years ago. This culture further expanded on fine bone tool work, including barbed spearheads and fishhooks (Figure 12.16).<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"511\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image2-1-1.jpg\" alt=\"Long, thin spear tips. Many have barbs, others are smooth.\" width=\"511\" height=\"494\" \/> Figure 12.16: This drawing from 1891 shows an array of Magdalenian-style barbed points found in the burial of a reindeer hunter. They were carved from antler. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:La_station_quaternaire_de_Raymonden_(...)Hardy_Michel_bpt6k5567846s_(2).jpg\">La station quaternaire de Raymonden (...)Hardy Michel bpt6k5567846s (2)<\/a> by M. F\u00e9auxis, original by Michel Hardy (1891), is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.[\/caption]\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Among the many European sites dating to the Later Stone Age, the famous cave art sites deserve mention. Chauvet-Pont-d'Arc Cave in southern France dates to separate Aurignacian occupations 31,000 years ago and 26,000 years ago. Over a hundred art pieces representing 13 animal species are preserved, from commonly depicted deer and horses to rarer rhinos and owls. Another French cave with art is Lascaux, which is several thousand years younger at 17,000 years ago in the Magdalenian period. At this site, there are over 6,000 painted figures on the walls and ceiling (Figure 12.17). Scaffolding and lighting must have been used to make the paintings on the walls and ceiling deep in the cave. Overall, visiting Lascaux as a contemporary must have been an awesome experience: trekking deeper in the cave lit only by torches giving glimpses of animals all around as mysterious sounds echoed through the galleries.<\/p>\r\n&nbsp;\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"605\"]<img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image24-2-1.jpg\" alt=\"Charcoal painting of a bull seen from the side.\" width=\"605\" height=\"454\" \/> Figure 12.17: Photograph of just one surface with cave art at Lascaux Cave. The most prominent piece here is the Second Bull, found in a chamber called the Hall of Bulls. Smaller cattle and horses are also visible. Credit: <a href=\"https:\/\/whc.unesco.org\/en\/documents\/108435\">Lascaux cave (document 108435) Prehitoric Sites and Decorated Caves of the V\u00e9z\u00e8re Valley (France)<\/a> by Francesco Bandarin, <a href=\"https:\/\/whc.unesco.org\/\">\u00a9 UNESCO<\/a>, has been modified (color modified) and is under a <a href=\"https:\/\/whc.unesco.org\/en\/licenses\/6\">CC BY-SA 3.0 License<\/a>.[\/caption]\r\n<h4 class=\"import-Normal\"><span style=\"background-color: #ff99cc\"><em>Summary of Modern <\/em>H. sapiens<em> in Europe<\/em><\/span><\/h4>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">Study of Europe in the Upper Paleolithic gives a more detailed view of the general pattern of biological and cultural change linked with the arrival of modern <em>Homo sapiens<\/em>. The modern humans experienced a rapidly changing culture that went through waves of complexity and refinement. Skeletally, the increasing globularity of the cranium and the gracility of the rest of the skeleton continued, though with unique regional traits, too. The cave art sites showed a deeper exploration of creativity though the exact meaning is unclear. With survival dependent on the surrounding ecology, painting the figures may have connected people to important and impressive wildlife at both a physical and spiritual level. Both reverence for animals and the use of caves for an enhanced sensory experience are common to cultures past and present.<\/span><\/p>\r\n\r\n<h3 class=\"import-Normal\"><strong>Peopling of the Americas<\/strong><\/h3>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">By 25,000 years ago, our species was the only member of <em>Homo<\/em> left on Earth. Gone were the Neanderthals, Denisovans, <em>Homo naledi,<\/em> and <em>Homo floresiensis<\/em>. The range of modern <em>Homo sapiens<\/em> kept expanding eastward into\u2014using the name given to this area by Europeans much later\u2014the Western Hemisphere. This section will address what we know about the peopling of the Americas, from the first entry to these continents to the rapid spread of Indigenous Americans across its varied environments.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">While evidence points to an ancient land bridge called [pb_glossary id=\"1778\"]<strong>Beringia<\/strong>[\/pb_glossary] that allowed people to cross from what is now northeastern Siberia into modern-day Alaska, what people did to cross this land bridge is still being investigated. For most of the 20th century, the accepted theory was the [pb_glossary id=\"1779\"]<strong>Ice-Free Corridor model<\/strong>[\/pb_glossary]. It stated that northeast Asians (East Asians and Siberians) first expanded across Beringia inland through a passage between glaciers that opened into the western Great Plains of the United States, just east of the Rocky Mountains, around 13,000 years ago (Swisher et al. 2013). While life up north in the cold environment would have been harsh, migrating birds and an emerging forest might have provided sustenance as generations expanded through this land (Potter et al. 2018).<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">However, in recent decades, researchers have accumulated evidence against the Ice-Free Corridor model. Archaeologist K. R. Fladmark (1979) brought the alternate [pb_glossary id=\"1780\"]<strong>Coastal Route model<\/strong>[\/pb_glossary] into the archaeological spotlight; researcher Jon M. Erlandson has been at the forefront of compiling support for this theory (Erlandson et al. 2015). The new focus is the southern edge of the land bridge instead of its center: About 16,000 years ago, members of our species expanded along the coastline from northeast Asia, east through Beringia, and south down the Pacific Coast of North America while the inland was still sealed off by ice. The coast would have been free of ice at least part of the year, and many resources would have been found there, such as fish (e.g., salmon), mammals (e.g., whales, seals, and otters), and plants (e.g., seaweed).<\/p>\r\n\r\n<h4 class=\"import-Normal\"><em>South through the Americas<\/em><\/h4>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">When the first modern <em>Homo sapiens<\/em> reached the Western Hemisphere, the spread through the Americas was rapid. Multiple migration waves crossed from North to South America (Posth et al. 2018). Our species took advantage of the lack of hominin competition and the bountiful resources both along the coasts and inland. The Americas had their own wide array of megafauna, which included woolly mammoths (Figure 12.18), mastodons, camels, horses, ground sloths, giant tortoises, and\u2014a favorite of researchers\u2014a two-meter-tall beaver. The reason we cannot see these amazing animals today may be that resources gained from these fauna were crucial to the survival for people over 12,000 years ago (Araujo et al. 2017). Several sites are notable for what they add to our understanding of the distant past in the Americas, including interactions with megafauna and other elements of the environment.<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"alignleft\" width=\"242\"]<img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image3-2-1.jpg\" alt=\"A mammoth model with long curving tusks.\" width=\"242\" height=\"323\" \/> Figure 12.18: Life-size reconstruction of a woolly mammoth at the Page Museum, part of the La Brea Tar Pits complex in Los Angeles, California. Outside of Africa, megafauna such as this went extinct around the time that humans entered their range. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-14\/\">Woolly Mammoth<\/a> (at <a href=\"https:\/\/tarpits.org\/\">La Brea Tar Pits &amp; Museum<\/a>) by Keith Chan is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.[\/caption]\r\n<p class=\"import-Normal\"><span style=\"background-color: #ff9900\">A 2019 discovery may allow researchers to improve theories about the peopling of the Americas. In White Sands National Park, New Mexico, 60 human footprints have been astonishingly dated to around 22,000 years ago (Bennett et al. 2021). This date and location do not match either the Ice-Free Corridor or Coastal Route models. Researchers are now working to verify the find and adjust previous models to account for the new evidence. This groundbreaking find is sparking new theories; it is another example of the fast pace of research performed on our past.<\/span><\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Monte Verde is a landmark site that shows that the human population had expanded down the whole vertical stretch of the Americas to Chile by 14,600 years ago, <span style=\"background-color: #ffff00\">only a few thousand years after humans first entered the Western Hemisphere from Alaska.<\/span> The site has been excavated by archaeologist Tom D. Dillehay and his team (2015). The remains of nine distinct edible species of seaweed at the site shows familiarity with coastal resources and relates to the Coastal Route model by showing a connection between the inland people and the sea.<\/p>\r\n\r\n[caption id=\"\" align=\"alignright\" width=\"254\"]<img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image21-4.png\" alt=\"A long stone point with small chips around the edge.\" width=\"254\" height=\"362\" \/> Figure 12.19: The Clovis point has a distinctive structure. It has a wide tip, and its base has two small projections. This example was carved from chert and found in north-central Ohio, dated to around 11,000 years ago. Credit: <a href=\"https:\/\/www.si.edu\/object\/chndm_15.2012.25\">Clovis Point<\/a> (15.2012.25) by <a href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [Department of Anthropology; Cooper Hewitt, Smithsonian Design Museum] <a href=\"https:\/\/www.si.edu\/termsofuse\">is used for educational and non-commercial purposes as outlined by the Smithsonian.<\/a>[\/caption]\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Named after the town in New Mexico, the Clovis stone-tool style is the first example of a widespread culture across much of North America, between 13,400 and 12,700 years ago (Miller, Holliday, and Bright 2013). Clovis points were fluted with two small projections, one on each end of the base, facing away from the head (Figure 12.19). The stone points found at this site match those found as far as the Canadian border and northern Mexico, and from the west coast to the east coast of the United States. Fourteen Clovis sites also contained the remains of mammoths or mastodons, suggesting that hunting megafauna with these points was an important part of life for the Clovis people. After the spread of the Clovis style, it diversified into several regional styles, keeping some of the Clovis form but also developing their own unique touches.<\/p>\r\n<span style=\"text-decoration: underline;background-color: #ff99cc\">(maybe inlcude a special topic\/dig deeper from Dr.Steeves talking about Clovis culture and effects on Indigenous histories)<\/span>\r\n<h4 class=\"import-Normal\"><span style=\"background-color: #ff99cc\"><em>Summary of Modern <\/em>H. sapiens<em> in the Americas<\/em><\/span><\/h4>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">Research in Native American origins found some surprising details, refining older models. Genetically, the migration can be considered one long period of movement with splits into regional populations. This finding matches the sudden appearance and diversification of the homegrown Clovis culture. A few thousand years after arrival into the hemisphere, people had already covered the Americas from north to south.<\/span><\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">The peopling of the Americas also had a lot of elements in common with the prior spread of humans across Africa, Europe, Asia, and Australia. In all of these expansions, these pioneers explored new lands that tested their ability to adapt, both culturally and biologically. Besides stone-tool technology, the use of ochre as decoration was seen from South Africa to South America. The coasts and rivers were likely avenues in the movement of people, artifacts, and ideas, outlining the land masses while providing access to varied environments. The presence of megafauna aided human success, but this resource was eventually depleted in many parts of the world.<\/span><\/p>\r\n\r\n<h3 class=\"import-Normal\"><strong>The Big Picture: The Assimilation Hypothesis<\/strong><\/h3>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">How do researchers make sense of all of these modern <em>Homo sapiens<\/em> discoveries that cover over 300,000 years of time and stretch across every continent except Antarctica? How was modern <em>Homo sapiens<\/em> related to archaic <em>Homo sapiens<\/em>?<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The <strong>[pb_glossary id=\"1782\"]Assimilation hypothesis[\/pb_glossary]<\/strong> proposes that modern <em>Homo sapiens<\/em> evolved in Africa first and expanded out but also interbred with the archaic <em>Homo sapiens<\/em> they encountered outside Africa (Figure 12.20). This hypothesis is powerful since it explains why Africa has the oldest modern human fossils, why early modern humans found in Europe and Asia bear a resemblance to the regional archaics, and why traces of archaic DNA can be found in our genomes today (Dannemann and Racimo 2018; Reich et al. 2010; Reich et al. 2011; Slatkin and Racimo 2016; Smith et al. 2017; Wall and Yoshihara Caldeira Brandt 2016).<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"443\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image28-2.png\" alt=\"African Homo erectus expands and gives rise to archaics and modern Homo sapiens groups.\" width=\"443\" height=\"471\" \/> Figure 12.20: This diagram shows archaic humans, having evolved from Homo erectus, expanded from Africa and established the Neanderthal and Denisovan groups. In Africa, archaic humans evolved modern traits and expanded from the continent as well, interbreeding with two archaic groups across Europe and Asia. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-14\/\">Assimilation Model (Figure 12.23)l<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Keith Chan and Katie Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.[\/caption]\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">While researchers have produced a model that satisfies the data, there are still a lot of questions for paleoanthropologists to answer regarding our origins. What were the patterns of migration in each part of the world? Why did the archaic humans go extinct? In what ways did archaic and modern humans interact? The definitive explanation of how our species started and what our ancestors did is still out there to be found. You are now in a great place to welcome the next discovery about our distant past\u2014maybe you\u2019ll even contribute to our understanding as well.<\/p>\r\n\r\n<h2 class=\"import-Normal\">The Chain Reaction of Agriculture<\/h2>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ccffcc\">While it may be hard to imagine today, for most of our species\u2019 existence we were nomadic: moving through the landscape without a singular home. Instead of a refrigerator or pantry stocked with food, we procured nutrition and other resources as needed based on what was available in the environment.<\/span> <span style=\"background-color: #ffff00\">Instead of collecting and displaying shelves of stuff, we kept our possessions small for mobility.<\/span> <span style=\"background-color: #ff99cc\">This section gives an overview of how the foraging lifestyle enabled the expansion of our species and how the invention of a new way of life caused a chain reaction of cultural change.<\/span><\/p>\r\n\r\n<h3 class=\"import-Normal\"><strong>The Foraging Tradition<\/strong><\/h3>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">There are a variety of possible <strong>[pb_glossary id=\"1783\"]subsistence strategies[\/pb_glossary]<\/strong>, or methods of finding sustenance and resources. To understand our species is to understand the subsistence strategy of <strong>[pb_glossary id=\"1114\"]foraging[\/pb_glossary]<\/strong>, or the search for resources in the environment. While most (but not all) humans today live in cultures that practice <strong>[pb_glossary id=\"1784\"]agriculture[\/pb_glossary] <\/strong>(whereby we greatly shape the environment to mass produce what we need), we have spent far more time as nomadic foragers than as settled agriculturalists. As such, <span style=\"background-color: #ffff00\">our traits have evolved to be primarily geared toward foraging. For instance, our efficient bipedalism allows persistence-hunting across long distances as well as movement from resource to resource.<\/span><\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">How does human foraging, also known as hunting and gathering, work? Anthropologists have used all four fields to answer this question (see Ember n.d.). Typically, people formed [pb_glossary id=\"1785\"]<strong>bands<\/strong>[\/pb_glossary], or kin-based groups of around 50 people or less (rarely over 100). A band\u2019s organization would be [pb_glossary id=\"1786\"]<strong>e<\/strong><strong>galitarian<\/strong>[\/pb_glossary], with a flexible hierarchy based on an individual\u2019s age, level of experience, and relationship with others. Everyone would have a general knowledge of the skills assigned to their gender roles, rather than specializing in different occupations. A band would be able to move from place to place in the environment, using knowledge of the area to forage (Figure 12.21). In varied environments\u2014from savannas to tropical forests, deserts, coasts, and the Arctic circle\u2014people found sustenance needed for survival. <span style=\"background-color: #ffff00\">Our species\u2019s omnivorousness and cultural abilities led us to excel in the generalist-specialist niche.<\/span><\/p>\r\n&nbsp;\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"565\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image22.jpg\" alt=\"A hunter holding a bow is crouched among dry grass.\" width=\"565\" height=\"377\" \/> Figure 12.21: A present-day San man in Namibia demonstrates hunting using archery. Anthropologists study the San today to learn about the persistence of foraging as a viable lifestyle, while noting how these cultures have changed over time and how they interact with other groups. Credit: <a href=\"https:\/\/www.flickr.com\/photos\/charlesfred\/2129551464\">San hunter w\u0131th bow and arrow<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/charlesfred\/\">CharlesFred<\/a> has been modified (color modified) and is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License.<\/a>[\/caption]\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ffff00\">Humans made extensive use of the foraging subsistence strategy, but this lifestyle did have limitations. The ease of foraging depended on the richness of the environment. Due to the lack of storage, resources had to be dependably found when needed. While a bountiful environment would require just a few hours of foraging a day and could lead to a focus on one location, the level and duration of labor increased greatly in poor or unreliable environments.<\/span> Labor was also needed to process the acquired resources, which contributed to the foragers\u2019 daily schedule (Crittenden and Schnorr 2017).<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ffff00\">The adaptations to foraging found in modern <em>Homo sapiens<\/em> may explain why our species became so successful both within Africa and in the rapid expansion around the world. Overcoming the limitations, each generation at the edge of our species\u2019s range would have found it beneficial to expand a little further, keeping contact with other bands but moving into unexplored territory where resources were more plentiful. The cumulative effect would have been the spread of modern <em>Homo sapiens<\/em> across continents and hemispheres.<\/span><\/p>\r\n\r\n<h2 class=\"import-Normal\"><strong>Why Agriculture?<\/strong><\/h2>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">After hundreds of thousands of years of foraging, some groups of people around 12,000 years ago started to practice agriculture. This transition, called the [pb_glossary id=\"1787\"]<strong>Neolithic Revolution<\/strong>,[\/pb_glossary] occurred at the start of the <strong>Holocene<\/strong> epoch. While the reasons for this global change are still being investigated, two likely co-occurring causes are a growing human population and natural global climate change.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Overcrowding could have affected the success of foraging in the environment, leading to the development of a more productive subsistence strategy (Cohen 1977). Foraging works best with low population densities since each band needs a lot of space to support itself. If too many people occupy the same environment, they deplete the area faster. The high population could exceed the <strong>[pb_glossary id=\"1788\"]carrying capacity[\/pb_glossary]<\/strong>, or number of people a location can reliably support. Reaching carrying capacity on a global level due to growing population and limited areas of expansion would have been an increasingly pressing issue after the expansion through the major continents by 14,600 years ago.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">A changing global climate immediately preceded the transition to agriculture, so researchers have also explored a connection between the two events. Since the <strong>[pb_glossary id=\"1789\"]Last Glacial Maximum[\/pb_glossary]<\/strong> of 23,000 years ago, the Earth slowly warmed. Then, from 13,000 to 11,700 years ago, the temperature in most of the Northern Hemisphere dropped suddenly in a phenomenon called the [pb_glossary id=\"1790\"]<strong>Younger Dryas<\/strong>.[\/pb_glossary] Glaciers returned in Europe, Asia, and North America. In Mesopotamia, which includes the Levant, the climate changed from warm and humid to cool and dry. The change would have occurred over decades, disrupting the usual nomadic patterns and subsistence of foragers around the world. The disruption to foragers due to the temperature shift could have been a factor in spurring a transition to agriculture. Researchers Gregory K. Dow and colleagues (2009) believe that foraging bands would have clustered in the new resource-rich places where people started to direct their labor to farming the limited area. After the Younger Dryas ended, people expanded out of the clusters with their agricultural knowledge (Figure 12.22).<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"570\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image7-6.png\" alt=\"Map shows that agriculture was invented in at least six parts of the world.\" width=\"570\" height=\"267\" \/> Figure 12.22: The map shows the areas where agriculture was independently invented around the world and where they spread. Blue arrows show the spread of agriculture from these zones to other regions. <a href=\"https:\/\/docs.google.com\/document\/d\/1VUDKMBJYS_jNONjLxT04jQN0_z9Ua50BRN6auGSHUuU\/edit\">A full text description of this image is available<\/a>. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Centres_of_origin_and_spread_of_agriculture.svg\">Centres of origin and spread of agriculture<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Joe_Roe\">Joe Roe<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.[\/caption]\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The double threat of the limitation of human continental expansion and the sudden global climate change may have placed bands in peril as more populations outpaced their environment\u2019s carrying capacity. <span style=\"background-color: #ffff00\">Not only had a growing population led to increased competition with other bands, but environments worldwide shifted to create more uncertainty. As people in different areas around the world faced this unpredictable situation, they became the independent inventors of agriculture.<\/span><\/p>\r\n\r\n<h2 class=\"import-Normal\"><strong>Agriculture around the World<\/strong><\/h2>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Due to global changes to the human experience starting from 12,000 years ago, <span style=\"background-color: #ffff00\">cultures with no knowledge of each other turned toward intensely farming their local resources<\/span> (see Figure 12.22). <span style=\"background-color: #ffff00\">The first farmers engaged in artificial selection of their domesticates to enhance useful traits over generations. The switch to agriculture took time and effort with no guarantee of success and constant challenges (e.g. fires, droughts, diseases, and pests).<\/span> The regions with the most widespread impact in the face of these obstacles became the primary centers of agriculture (Figure 12.23; Fuller 2010):<\/p>\r\n\r\n<ul>\r\n \t<li class=\"import-Normal\" style=\"text-indent: 0pt\">Mesopotamia: The Fertile Crescent from the Tigris and Euphrates rivers through the Levant was where bands started to domesticate plants and animals around 12,000 years ago. The connection between the development of agriculture and the Younger Dryas was especially strong here. Farmed crops included wheat, barley, peas, and lentils. This was also where cattle, pigs, sheep, and goats were domesticated.<\/li>\r\n \t<li class=\"import-Normal\" style=\"text-indent: 0pt\">South and East Asia: Multiple regions across this land had varieties of rice, millet, and soybeans by 10,000 years ago. Pigs were farmed with no connection to Mesopotamia. Chickens were also originally from this region, bred for fighting first and food second.<\/li>\r\n \t<li class=\"import-Normal\" style=\"text-indent: 0pt\">New Guinea: Agriculture started here 10,000 years ago. Bananas, sugarcane, and taro were native to this island. Sweet potatoes were brought back from voyages to South America around the year C.E. 1000. No known animal farming occurred here.<\/li>\r\n \t<li class=\"import-Normal\" style=\"text-indent: 0pt\">Mesoamerica: Agriculture from Central Mexico to northern South America also occurred from 10,000 years ago; it was also only plant based. Maize was a crop bred from teosinte grass, which has become one of the global staples. Beans, squash, and avocados were also grown in this region.<\/li>\r\n \t<li class=\"import-Normal\" style=\"text-indent: 0pt\">The Andes: Starting around 8,000 years ago, local domesticated plants started with squash but later included potatoes, tomatoes, beans, and quinoa. Maize was brought down from Mesoamerica. The main farm animals were llamas, alpacas, and guinea pigs.<\/li>\r\n \t<li class=\"import-Normal\" style=\"text-indent: 0pt\">Sub-Saharan Africa: This region went through a change 5,000 years ago called the Bantu expansion. The Bantu agriculturalists were established in West Central Africa and then expanded south and east. Native varieties of rice, yams, millet, and sorghum were grown across this area. Cattle were also domesticated here.<\/li>\r\n \t<li class=\"import-Normal\" style=\"text-indent: 0pt\">Eastern North America: This region was the last major independent agriculture center, from 4,000 years ago. Squash and sunflower are the produce from this region that are most known today, though sumpweed and pitseed goosefoot were also farmed. Hunting was still the main source of animal products.<\/li>\r\n<\/ul>\r\n[caption id=\"\" align=\"aligncenter\" width=\"482\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image23-1-1.jpg\" alt=\"Farmers plow a flooded field. Each plow is pulled by two oxen. \" width=\"482\" height=\"320\" \/> Figure 12.23: Rice farmers in the present day using draft cattle to prepare their field. Credit: <a href=\"https:\/\/www.flickr.com\/photos\/ricephotos\/7554483250\">Plowing muddy field using cattle<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/ricephotos\/\">IRRI Photos<\/a> (International Rice Research Institute) has been modified (color modified) and is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a>.[\/caption]\r\n<p class=\"import-Normal\">By 5,000 years ago, our species was well within the Neolithic Revolution. Agriculturalists spread to neighboring parts of the world with their domesticates, further expanding the use of this subsistence strategy. <span style=\"background-color: #ffff00\">From this point, the human species changed from being primarily foragers to primarily agriculturalists with skilled control of their environments.<\/span> The planet changed from mostly unaffected by human presence to being greatly transformed by humans. The revolution took millennia, but it was a true revolution as our species\u2019 lifestyle was dramatically reshaped.<\/p>\r\n\r\n<h3 class=\"import-Normal\"><strong>Cultural Effects of Agriculture<\/strong><\/h3>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The worldwide adoption of agriculture altered the course of human culture and history forever. The core change in human culture due to agriculture is the move toward not moving: rather than live a nomadic lifestyle, farmers had to remain in one area to tend to their crops and livestock. The term for living bound to a certain location is <strong>[pb_glossary id=\"1793\"]sedentarism[\/pb_glossary]<\/strong>. <span style=\"background-color: #ffff00\">This led to new aspects of life that were uncommon among foragers: the construction of permanent shelters and agricultural infrastructure, such as fields and irrigation, plus the development of storage technology, such as pottery, to preserve extra resources in case of future instability.<\/span><\/p>\r\n\r\n\r\n[caption id=\"\" align=\"alignright\" width=\"359\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image20-1-1.jpg\" alt=\"Multistory buildings surrounding a greek-style plaza.\" width=\"359\" height=\"270\" \/> Figure 12.24: View of downtown San Diego taken by the author at a shopping complex during a break from jury duty. Here, people live amongst structures that facilitate commerce, government, tourism, and art. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-14\/\">Downtown San Diego (October 13, 2016; Figure 12.28)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Keith Chan is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>.[\/caption]\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The high productivity of successful agriculture sparked further changes (Smith 2009). <span style=\"background-color: #ffff00\">Since successful agriculture produced a much-greater amount of food and other resources per unit of land compared to foraging, the population growth rate skyrocketed.<\/span> <span style=\"background-color: #ccffcc\">The surplus of a bountiful harvest also provided insurance for harder times, reducing the risk of famine. Changes happened to society as well. With a few farming households producing enough food to feed many others, other people could focus on other tasks. So began specialization into different occupations such as craftspeople, traders, religious figures, and artists, spurring innovation in these areas as people could now devote time and effort toward specific skills. These interdependent people would settle an area together for convenience. The growth of these settlements led to <strong>[pb_glossary id=\"1794\"]urbanization[\/pb_glossary]<\/strong>, the founding of cities that became the foci of human interaction (Figure 12.24).<\/span><\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The formation of cities led to new issues that sparked the growth of further specializations, called <strong>[pb_glossary id=\"1795\"]institutions[\/pb_glossary]<\/strong>. These are cultural constructs that exist beyond the individual and have wide control over a population. Leadership of these cities became hierarchical with different levels of rank and control. The stratification of society increased social inequality between those with more or less power over others. Under leadership, people built impressive <strong>[pb_glossary id=\"1796\"]monumental architecture[\/pb_glossary]<\/strong>, such as pyramids and palaces, that embodied the wealth and power of these early cities. Alliances could unite cities, forming the earliest states. In several regions of the world, state organization expanded into empires, wide-ranging political entities that covered a variety of cultures.<\/p>\r\n<span style=\"text-decoration: underline;background-color: #00ffff\">(Inlcude Special Topic about the Haudesaunee\/Iroquois confederacy)<\/span>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Urbanization brought new challenges as well. The concentration of sedentary peoples was ideal for infectious diseases to thrive since they could jump from person to person and even from livestock to person (Armelagos, Brown, and Turner 2005). While successful agriculture provided a large surplus of food to thwart famine, the food produced offered less diverse food sources than foragers\u2019 diets (Cohen and Armelagos 1984; Cohen and Crane-Kramer 2007). This shift in nutrition caused other diseases to flourish among those who adopted farming, such as dental cavities and malocclusion (the misalignment of teeth caused by soft, agricultural diets). The need to extract \u201cwisdom teeth\u201d or third molars seen in agricultural cultures today stems from this misalignment between the environment our ancestors adapted to and our lifestyles today.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">As the new disease trends show, the adoption of agriculture and the ensuing cultural changes were not entirely positive. It is also important to note that this is not an absolutely linear progression of human culture from simple to complex. In many cases, empires have collapsed and, in some cases, cities dispersed to low-density bands that rejected institutions. However, a global trend has emerged since the adoption of agriculture, wherein population and social inequality have increased, leading to the massive and influential nation-states of today.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The rise of states in Europe has a direct impact on many of this book\u2019s topics. Science started as a European cultural practice by the upper class that became a standardized way to study the world. Education became an institution to provide a standardized path toward producing and gaining knowledge. The scientific study of human diversity, embroiled in the race concept that still haunts us today, was connected to the European slave trade and colonialism.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Also starting in Europe, the Industrial Revolution of the 19th century turned cities into centers of mass manufacturing and spurred the rapid development of inventions (Figure 12.25). In the technologically interconnected world of today, human society has reached a new level of complexity with <strong>globalization<\/strong>. In this system, goods are mass-produced and consumed in different parts of the world, weakening the reliance on local farms and factories. The imbalanced relationship between consumers and producers of goods further increases economic inequality.<\/p>\r\n&nbsp;\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"465\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image1-3.jpg\" alt=\"A yellow farm vehicle driving into crops in a field.\" width=\"465\" height=\"310\" \/> Figure 12.25: This combine harvester can collect and process grain at a massive scale. Our food now commonly comes from enormous farms located around the world. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Combine_CR9060.jpeg\">Combine CR9060<\/a> by Hertzsprung is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.[\/caption]\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">As states based on agriculture and industry keep exerting influence on humanity today, there are people, like the Hadzabe of Tanzania, who continue to live a lifestyle centered on foraging. Due to the overwhelming force that agricultural societies exert, foragers today have been marginalized to live in the least habitable parts of the world\u2014the areas that are not conducive to farming, such as tropical rainforests, deserts, and the Arctic (Headland et al. 1989). Foragers can no longer live in the abundant environments that humans would have enjoyed before the Neolithic Revolution. Interactions with agriculturalists are typically imbalanced, with trade and other exchanges heavily favoring the larger group. One of anthropology\u2019s important roles today is to intelligently and humanely manage equitable interactions between people of different backgrounds and levels of influence.<\/p>\r\n\r\n<div class=\"textbox\">\r\n<h2 class=\"import-Normal\">Special Topic: Indigenous Land Management<\/h2>\r\n<p class=\"import-Normal\">Insight into the lives of past modern humans has evolved as researchers revise previous theories and establish new connections with Indigenous knowledge holders.<\/p>\r\n<p class=\"import-Normal\">The outdated view of foraging held that people lived off of the land without leaving an impact on the environment. Accompanying this idea was anthropologist Marshall Sahlins\u2019s (1968) proposal that foragers were the \u201coriginal affluent society\u201d since they were meeting basic needs and achieving satisfaction with less work hours than agriculturalists and city-dwellers. This view countered an earlier idea that foragers were always on the brink of starvation. Sahlins\u2019s theory took hold in the public eye as an attractive counterpoint to our busy contemporary lives in which we strive to meet our endless wants.<\/p>\r\n<p class=\"import-Normal\">A fruitful type of study involving researchers collaborating with Indigenous experts has found that foragers did not just live off the land with minimal effort nor were they barely surviving in unchanging environments. Instead, they shaped the landscape to their needs using labor and strategies that were more subtle than what European colonizers and subsequent researchers were used to seeing. Research from two regions shows the latest developments in understanding Indigenous land management.<\/p>\r\n<p class=\"import-Normal\">In British Columbia, Canada, the bridging of scientific and Indigenous perspectives has shown that the forests of the region are not untouched wilderness but, rather, have been crafted by Indigenous peoples thousands of years ago. Forest gardens adjacent to archaeological sites show higher plant diversity than unmanaged places even after 150 years (Armstrong et al. 2021). On the coast, 3,500-year-old archaeological sites are evidence of constructed clam gardens, according to Indigenous experts (Lepofsky et al. 2015). Another project, in consultation with Elders of the T\u2019exelc (William Lakes First Nation) in British Columbia, introduced researchers to explanations of how forests were managed before the practice was disrupted by European colonialism (Copes-Gerbitz et al. 2021). Careful management of controlled fires reduced the density of the forest to favor plants such as raspberries and allow easier movement through the landscape.<\/p>\r\n<p class=\"import-Normal\">Similarly, the study of landscapes in Australia, in consultation with Aboriginal Australians today, shows that areas previously considered wilderness by scientists were actually the result of controlling fauna and fires. The presence of grasslands with adjacent forests were purposely constructed to attract kangaroos for hunting (Gammage 2008). People also managed other animal and insect life, from emus to caterpillars. In Tasmania, a shift from productive grassland to wildfire-prone rainforest occurred after Aboriginal Australian land management was replaced by British colonial rule (Fletcher, Hall, and Alexander 2021). The site of Budj Bim of the Gunditjmara people has archaeological features of <strong>[pb_glossary id=\"1798\"]aquaculture[\/pb_glossary]<\/strong>, or the farming of fish, that date back 6,600 years (McNiven et al. 2012; McNiven et al. 2015). These examples show that Indigenous knowledge of how to manipulate the environment may be invaluable at the state level, such as by creating an Aboriginal ranger program to guide modern land management.<\/p>\r\n\r\n<\/div>\r\n<h2 class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Conclusion<\/span><\/h2>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">Modern <em>Homo sapiens<\/em> is the species that took the hominin lifestyle the furthest to become the only living member of that lineage. The largest factor that allowed us to persist while other hominins went extinct was likely our advanced ability to culturally adapt to a wide variety of environments. Our species, with its skeletal and behavioral traits, was well-suited to be generalist-specialists who successfully foraged across most of the world\u2019s environments. The biological basis of this adaptation was our reorganized brain that facilitated innovation in cultural adaptations and intelligence for leveraging our social ties and finding ways to acquire resources from the environment. As the brain\u2019s ability increased, it shaped the skull by reducing the evolutionary pressure to have large teeth and robust cranial bones to produce the modern <em>Homo sapiens<\/em> face.<\/span><\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">Our ability to be generalist-specialists is seen in the geographical range that modern <em>Homo sapiens<\/em> covered in 300,000 years. In Africa, our species formed from multiregional gene flow that loosely connected archaic humans across the continent. People then expanded out to the rest of the continental Eurasia and even further to the Americas.<\/span><\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">For most of our species\u2019s existence, foraging was the general subsistence strategy within which people specialized to culturally adapt to their local environment. With omnivorousness and mobility, people found ways to extract and process resources, shaping the environment in return. When resource uncertainty hit the species, people around the world focused on agriculture to have a firmer control of sustenance. The new strategy shifted human history toward exponential growth and innovation, leading to our high dependence on cultural adaptations today.<\/span><\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">While a cohesive image of our species has formed in recent years, there is still much to learn about our past. The work of many driven researchers shows that there are amazing new discoveries made all the time that refine our knowledge of human evolution. Technological innovations such as DNA analysis enable scientists to approach lingering questions from new angles. The answers we get allow us to ask even more insightful questions that will lead us to the next revelation. Like the pink limestone strata at Jebel Irhoud, previous effort has taken us so far and you are now ready to see what the next layer of discovery holds.<\/span><\/p>\r\n\r\n<div class=\"textbox\">\r\n<h2 class=\"import-Normal\"><del>Special Topic:<\/del> The Future of Humanity<\/h2>\r\n<p class=\"import-Normal\">A common question stemming from understanding human evolution is: What will the genetic and biological traits of our species be hundreds of thousands of years in the future? When faced with this question, people tend to think of directional selection. Maybe our braincases will be even larger, resembling the large-headed and small-bodied aliens of science fiction (Figure 12.26). Or, our hands could be specialized for interacting with our touch-based technology with less risk of repetitive injury. These ideas do not stand up to scrutiny. Since natural selection is based on adaptations that increase reproductive success, any directional change must be due to a higher rate of producing successful offspring compared to other alleles. Larger brains and more agile fingers would be convenient to possess, but they do not translate into an increase in the underlying allele frequencies.<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"571\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image8-4.png\" alt=\"One human has typical features; the other has a tall braincase.\" width=\"571\" height=\"279\" \/> Figure 12.26: Will we evolve toward even more globular brains? Actually, this trend is not likely to continue for our species. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-14\/\">Hypothetical image of future human evolution (Figure 12.30)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.[\/caption]\r\n<p class=\"import-Normal\">Scientists are hesitant to professionally speculate on the unknowable, and we will never know what is in store for our species one thousand or one million years from now, but there are two trends in human evolution that may carry on into the future: increased genetic variation and a reduction in regional differences.<\/p>\r\n<p class=\"import-Normal\">Rather than a directional change, genetic variation in our species could expand. Our technology can protect us from extreme environments and pathogens, even if our biological traits are not tuned to handle these stressors. The rapid pace of technological advancement means that biological adaptations will become less and less relevant to reproductive success, so nonbeneficial genetic traits will be more likely to remain in the gene pool. Biological anthropologist Jay T. Stock (2008) views environmental stress as needing to defeat two layers of protection before affecting our genetics. The first layer is our cultural adaptations. Our technology and knowledge can reduce pressure on one\u2019s genotype to be \u201cjust right\u201d to pass to the next generation. The second defense is our flexible physiology, such as our acclimatory responses. Only stressors not handled by these powerful responses would then cause natural selection on our alleles. These shields are already substantial, and cultural adaptations will only keep increasing in strength.<\/p>\r\n<p class=\"import-Normal\">The increasing ability to travel far from one\u2019s home region means that there will be a mixing of genetic variation on a global level in the future of our species. In recent centuries, gene flow of people around the world has increased, creating admixture in populations that had been separated for tens of thousands of years. For skin color, this means that populations all around the world could exhibit the whole range of skin colors, rather than the current pattern of decreasing melanin pigment farther from the equator. The same trend of intermixing would apply to all other traits, such as blood types. While our genetics will become more varied, the variation will be more intermixed instead of regionally isolated.<\/p>\r\n<p class=\"import-Normal\">Our distant descendants will not likely be dextrous ultraintellectuals; more likely, they will be a highly variable and mobile species supported by novel cultural adaptations that make up for any inherited biological limitations. Technology may even enable the editing of DNA directly, changing these trends. With the uncertainty of our future, these are just the best-educated guesses for now. Our future is open and will be shaped little by little by the environment, our actions, and the actions of our descendants.<\/p>\r\n\r\n<\/div>\r\n<h2 class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Hominin Species Summary<\/span><\/h2>\r\n<div style=\"text-align: left\">\r\n<table class=\"aligncenter\" style=\"width: 450pt\">\r\n<tbody>\r\n<tr class=\"Table1-R\" style=\"height: 0\">\r\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Modern<em> Homo sapiens<\/em><\/span><\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table1-R\" style=\"height: 0\">\r\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">315,000 years ago to present<\/span><\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table1-R\" style=\"height: 0\">\r\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\"><span style=\"color: #000000\">Starting in Africa, then expanding around the world<\/span><\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table1-R\" style=\"height: 0\">\r\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">Cro-Magnon individuals, discovered 1868 in Dordogne, France. Otzi the Ice Man, discovered 1991 in the Alps between Austria and Italy. Kennewick man, discovered 1996 in Washington state.<\/span><\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table1-R\" style=\"height: 0\">\r\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">1400 cc average<\/span><\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table1-R\" style=\"height: 0\">\r\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Extremely small with short cusps.<\/span><\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table1-R\" style=\"height: 0\">\r\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">An extremely globular brain case and gracile features throughout the cranium. The mandibular symphysis forms a chin at the anterior-most point.<\/span><\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table1-R\" style=\"height: 0\">\r\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\"><span style=\"color: #000000\">Gracile skeleton adapted for efficient bipedal locomotion at the expense of the muscular strength of most other large primates.<\/span><\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table1-R\" style=\"height: 0\">\r\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\"><span style=\"color: #000000\">Extremely extensive and varied culture with many spoken and written languages. Art is ubiquitous. Technology is broad in complexity and impact on the environment.<\/span><\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table1-R\" style=\"height: 0\">\r\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\r\n<p class=\"import-Normal\"><span style=\"color: #000000\">The only living hominin. Chimpanzees and bonobos are the closest living relatives.<\/span><\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr>\r\n<td><\/td>\r\n<td><\/td>\r\n<\/tr>\r\n<\/tbody>\r\n<\/table>\r\n<\/div>\r\n<div class=\"textbox shaded\">\r\n<h2 class=\"import-Normal\">Review Questions<strong>\r\n<\/strong><\/h2>\r\n<ul>\r\n \t<li>What are the skeletal and behavioral traits that define modern <em>Homo sapiens<\/em>? What are the evolutionary explanations for its presence?<\/li>\r\n \t<li>What are some creative ways that researchers have learned about the past by studying fossils and artifacts?<\/li>\r\n \t<li>How do the discoveries mentioned in \u201cFirst Africa, Then the World\u201d fit the Assimilation model?<\/li>\r\n \t<li>What is foraging? What adaptations do we have for this subsistence strategy? Could you train to be a skilled forager?<\/li>\r\n \t<li>What are aspects of your life that come from dependence on agriculture and its cultural effects? Where did the ingredients of your favorite foods originate from?<\/li>\r\n<\/ul>\r\n<\/div>\r\n<\/div>\r\n<h2 class=\"__UNKNOWN__\">Key Terms<\/h2>\r\n<div class=\"__UNKNOWN__\">\r\n<p class=\"import-Normal\"><strong>African multiregionalism<\/strong>: The idea that modern <em>Homo sapiens<\/em> evolved as a complex web of small regional populations with sporadic gene flow among them.<\/p>\r\n<p class=\"import-Normal\"><strong>Agriculture<\/strong>: The mass production of resources through farming and domestication.<\/p>\r\n<p class=\"import-Normal\"><strong>Aquaculture<\/strong>: The farming of fish using techniques such as trapping, channels, and artificial ponds.<\/p>\r\n<p class=\"import-Normal\"><strong>Assimilation <\/strong><strong>hypothesis<\/strong>: Current theory of modern human origins stating that the species evolved first in Africa and interbred with archaic humans of Europe and Asia.<\/p>\r\n<p class=\"import-Normal\"><strong>Atlatl<\/strong>: A handheld spear thrower that increased the force of thrown projectiles.<\/p>\r\n<p class=\"import-Normal\"><strong>Band<\/strong>: A small group of people living together as foragers.<\/p>\r\n<p class=\"import-Normal\"><strong>Beringia<\/strong>: Ancient landmass that connected Siberia and Alaska. The ancestors of Indigenous Americans would have crossed this area to reach the Americas.<\/p>\r\n<p class=\"import-Normal\"><strong>Carrying capacity<\/strong>: The amount of organisms that an environment can reliably support.<\/p>\r\n<p class=\"import-Normal\"><strong>Coastal Route model<\/strong>: Theory that the first Paleoindians crossed to the Americas by following the southern coast of Beringia.<\/p>\r\n<p class=\"import-Normal\"><strong>Early Modern <\/strong><strong><em>Homo sapiens<\/em><\/strong><strong>, Early Anatomically Modern Human<\/strong>: Terms used to refer to transitional fossils between archaic and modern <em>Homo sapiens<\/em> that have a mosaic of traits. Humans like ourselves, who mostly lack archaic traits, are referred to as Late Modern <em>Homo sapiens<\/em> and simply Anatomically Modern Humans.<\/p>\r\n<p class=\"import-Normal\"><strong>Egalitarian<\/strong>: Human organization without strict ranks. Foraging societies tend to be more egalitarian than those based on other subsistence strategies.<\/p>\r\n<p class=\"import-Normal\"><strong>Foraging<\/strong>: Lifestyle consisting of frequent movement through the landscape and acquiring resources with minimal storage capacity.<\/p>\r\n<p class=\"import-Normal\"><strong>Generalist-specialist niche<\/strong>: The ability to survive in a variety of environments by developing local expertise. Evolution toward this niche may have been what allowed modern <em>Homo sapiens<\/em> to expand past the geographical range of other human species.<\/p>\r\n<p class=\"import-Normal\"><strong>Globalization<\/strong>: A recent increase in the interconnectedness and interdependence of people that is facilitated with long-distance networks.<\/p>\r\n<p class=\"import-Normal\"><strong>Globular<\/strong>: Having a rounded appearance. Increased globularity of the braincase is a trait of modern <em>Homo sapiens<\/em>.<\/p>\r\n<p class=\"import-Normal\"><strong>Gracile<\/strong>: Having a smooth and slender quality; the opposite of robust.<\/p>\r\n<p class=\"import-Normal\"><strong>Holocene<\/strong>: The epoch of the Cenozoic Era starting around 12,000 years ago and lasting arguably through the present.<\/p>\r\n<p class=\"import-Normal\"><strong>Ice-Free Corridor model<\/strong>: Theory that the first Native Americans crossed to the Americas through a passage between glaciers.<\/p>\r\n<p class=\"import-Normal\"><strong>Institutions<\/strong>: Long-lasting and influential cultural constructs. Examples include government, organized religion, academia, and the economy.<\/p>\r\n<p class=\"import-Normal\"><strong>Last Glacial Maximum<\/strong>: The time 23,000 years ago when the most recent ice age was the most intense.<\/p>\r\n<p class=\"import-Normal\"><strong>Later Stone Age<\/strong>: Time period following the Middle Stone Age with a diversification in tool types, starting around 50,000 years ago.<\/p>\r\n<p class=\"import-Normal\"><strong>Levant<\/strong>: The eastern coast of the Mediterranean. The site of early modern human expansion from Africa and later one of the centers of agriculture.<\/p>\r\n<p class=\"import-Normal\"><strong>Megafauna<\/strong>: Large ancient animals that may have been hunted to extinction by people around the world.<\/p>\r\n<p class=\"import-Normal\"><strong>Mental eminence<\/strong>: The chin on the mandible of modern <em>H. sapiens<\/em>. One of the defining traits of our species.<\/p>\r\n<p class=\"import-Normal\"><strong>Microlith<\/strong>: Small stone tool found in the Later Stone Age; also called a bladelet.<\/p>\r\n<p class=\"import-Normal\"><strong>Middle Stone Age<\/strong>: Time period known for Mousterian lithics that connects African archaic to modern <em>Homo sapiens<\/em>.<\/p>\r\n<p class=\"import-Normal\"><strong>Monumental architecture<\/strong>: Large and labor-intensive constructions that signify the power of the elite in a sedentary society. A common type is the pyramid, a raised crafted structure topped with a point or platform.<\/p>\r\n<p class=\"import-Normal\"><strong>Mosaic<\/strong>: Composed from a mix or composite of traits.<\/p>\r\n<p class=\"import-Normal\"><strong>Neolithic Revolution<\/strong>: Time of rapid change to human cultures due to the invention of agriculture, starting around 12,000 years ago.<\/p>\r\n<p class=\"import-Normal\"><strong>Ochre<\/strong>: Iron-based mineral pigment that can be a variety of yellows, reds, and browns. Used by modern human cultures worldwide since at least 80,000 years ago.<\/p>\r\n<p class=\"import-Normal\"><strong>Sahul<\/strong>: Ancient landmass connecting New Guinea and Australia.<\/p>\r\n<p class=\"import-Normal\"><strong>Sedentarism<\/strong>: Lifestyle based on having a stable home area; the opposite of nomadism.<\/p>\r\n<p class=\"import-Normal\"><strong>Southern Dispersal model<\/strong>: Theory that modern <em>H. sapiens<\/em> expanded from East Africa by crossing the Red Sea and following the coast east across Asia.<\/p>\r\n<p class=\"import-Normal\"><strong>Subsistence strategy<\/strong>: The method an organism uses to find nourishment and other resources.<\/p>\r\n<p class=\"import-Normal\"><strong>Sunda<\/strong>: Ancient Asian landmass that incorporated modern Southeast Asia.<\/p>\r\n<p class=\"import-Normal\"><strong>Supraorbital torus<\/strong>: The bony brow ridge across the top of the eye orbits on many hominin crania.<\/p>\r\n<p class=\"import-Normal\"><strong>Upper Paleolithic<\/strong>: Time period considered synonymous with the Later Stone Age.<\/p>\r\n<p class=\"import-Normal\"><strong>Urbanization<\/strong>: The increase of population density as people settled together in cities.<\/p>\r\n<p class=\"import-Normal\"><strong>Wallacea<\/strong>: Archipelago southeast of Sunda with different biodiversity than Asia.<\/p>\r\n<p class=\"import-Normal\"><strong>Younger Dryas<\/strong>: The rapid change in global climate\u2014notably a cooling of the Northern Hemisphere\u201413,000 years ago.<\/p>\r\n\r\n<h2 class=\"import-Normal\">About the Author<strong>\r\n<\/strong><\/h2>\r\n<p class=\"import-Normal\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image14-3-1.jpg\" alt=\"A man with short, black hair stands outdoors with a setting sun behind him.\" width=\"282\" height=\"376\" \/><\/p>\r\n\r\n<h3 class=\"import-Normal\"><strong>Keith Chan, Ph.D.<\/strong><\/h3>\r\n<p class=\"import-Normal\">Grossmont-Cuyamaca Community College District and MiraCosta College, drkeithcchan@gmail.com, Dr. Keith Chan is an instructor of anthropology at Grossmont-Cuyamaca Community College District and MiraCosta College in San Diego County. He reached this step of his anthropological path after many memorable experiences across the country and the hemisphere. He earned a bachelor\u2019s degree in anthropology from the University of California, Berkeley, in 2001. As a graduate student at the University of Missouri, he traveled to Per\u00fa with teams of students to study skeletons in the archaeological record to understand the lives of ancient Andeans. He completed his dissertation and earned a Ph.D. in 2011. Inspired by many educators in his journey, Dr. Chan turned his career toward teaching anthropology and helping students understand and appreciate humanity.<\/p>\r\n\r\n<h2 class=\"import-Normal\">For Further Exploration<\/h2>\r\n<h3 class=\"import-Normal\" style=\"text-indent: 0pt\"><strong>Websites<\/strong><\/h3>\r\nFirst-person virtual tour of Lascaux cave with annotated cave art: Minist\u00e8re de la Culture and Mus\u00e9e d\u2019Arch\u00e9ologie Nationale. \u201c<a href=\"https:\/\/archeologie.culture.fr\/lascaux\/en\/visit-cave\" target=\"_blank\" rel=\"noopener\">Visit the cave<\/a>\u201d Lascaux website.\r\n\r\nOnline anthropology magazine articles related to paleoanthropology and human evolution: SAPIENS. \u201c<a href=\"https:\/\/www.sapies.org\/category\/evolution\/\" target=\"_blank\" rel=\"noopener\">Evolution<\/a>.\u201d <em>SAPIENS<\/em> website.\r\n\r\nVarious presentations of information about hominin evolution: Smithsonian Institution. \u201c<a href=\"https:\/\/humanorigins.si.edu\" target=\"_blank\" rel=\"noopener\">What does it mean to be human?<\/a>\u201d <em>Smithsonian National Museum of Natural History<\/em> website.\r\n\r\nMagazine-style articles on archaeology and paleoanthropology: ThoughtCo. \u201c<a href=\"https:\/\/www.thoughtco.com\/archaeology-4133504\" target=\"_blank\" rel=\"noopener\">Archaeology<\/a>.\u201d ThoughtCo. Website.\r\n\r\nDatabase of comparisons across hominins and primates: University of California, San Diego. \u201c<a href=\"https:\/\/carta.anthropogeny.org\/moca\/domains\" target=\"_blank\" rel=\"noopener\">MOCA Domains<\/a>.\u201d <em>Center for Academic Research &amp; Training in Anthropogeny<\/em> website.\r\n<h3><strong>Books<\/strong><\/h3>\r\nEngaging book that covers human-made changes to the environment with industrialization and globalization: Kolbert, Elizabeth. 2014. <em>The Sixth Extinction: An Unnatural History<\/em>. New York: Bloomsbury.\r\n\r\nOverview of what human life was like among the environmental shifts of the Ice Age: Woodward, Jamie. 2014. <em>The Ice Age: A Very Short Introduction<\/em>. Oxford: OUP Press.\r\n<h3><strong>Articles<\/strong><\/h3>\r\nRecent review paper about the current state of paleoanthropology research: Stringer, C. 2016. \u201c<a href=\"https:\/\/doi.org\/10.1098\/rstb.2015.0237\" target=\"_blank\" rel=\"noopener\">The Origin and Evolution of <em>Homo sapiens<\/em><\/a>.\u201d <em>Philosophical Transactions of the Royal Society B<\/em> 371 (1698).\r\n\r\nOverview of the history of American paleoanthropology and the many debates that have occurred over the years: Trinkaus, E. 2018. \u201cOne Hundred Years of Paleoanthropology: An American Perspective.\u201d <em>American Journal of Physical Anthropology<\/em> 165 (4): 638\u2013651.\r\n\r\nAmazing magazine article that synthesizes hominin evolution and why it is important to study this subject: Wheelwright, Jeff. 2015. \u201c<a href=\"https:\/\/discovermagazine.com\/2015\/may\/16-days-of-dysevolution\" target=\"_blank\" rel=\"noopener\">Days of Dysevolution<\/a>.\u201d <em>Discover<\/em> 36 (4): 33\u201339.\r\n\r\nFascinating research on \u00d6tzi, a mummy from 5,000 years ago: Wierer, Ursula, Simona Arrighi, Stefano Bertola, G\u00fcnther Kaufmann, Benno Baumgarten, Annaluisa Pedrotti, Patrizia Pernter, and Jacques Pelegrin. 2018. \u201cThe Iceman\u2019s Lithic Toolkit: Raw Material, Technology, Typology and Use.\u201d <em>PLOS One<\/em> 13 (6): e0198292. https:\/\/doi.org\/10.1371\/journal.pone.0198292.\r\n<h3><strong>Documentaries<\/strong><\/h3>\r\nPBS NOVA series covering the expansion of modern <em>Homo sapiens<\/em> and interbreeding with archaic humans: Brown, Nicholas, dir. 2015. <em>First Peoples<\/em>. Edmonton: Wall to Wall Television. Amazon Prime Video.\r\n\r\nPBS NOVA special featuring the footprints found in White Sands National Park: Falk, Bella, dir. 2016. <em>Ice Age Footprints<\/em>. Boston: Windfall Films. https:\/\/www.pbs.org\/wgbh\/nova\/video\/ice-age-footprints\/.\r\n\r\nPBS NOVA special about how modern humans evolved adaptations to different environments. Shows how present-day people live around the world: Thompson, Niobe, dir. 2016. <em>Great Human Odyssey<\/em>. Edmonton: Clearwater Documentary. <a class=\"rId132\" href=\"https:\/\/www.pbs.org\/wgbh\/nova\/evolution\/great-human-odyssey.html\">https:\/\/www.pbs.org\/wgbh\/nova\/evolution\/great-human-odyssey.html<\/a>.\r\n\r\n<\/div>\r\n<h2 class=\"__UNKNOWN__\">References<\/h2>\r\n<div class=\"__UNKNOWN__\">\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Araujo, Bernardo B. A., Luiz Gustavo R. Oliveira-Santos, Matheus S. Lima-Ribeiro, Jos\u00e9 Alexandre F. Diniz-Filho, and Fernando A. S. Fernandez. 2017. \u201cBigger Kill Than Chill: The Uneven Roles of Humans and Climate on Late Quaternary Megafaunal Extinctions.\u201d <em>Quaternary International<\/em> 431: 216\u2013222.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Armelagos, George J., Peter J. Brown, and Bethany Turner. 2005. \u201cEvolutionary, Historical, and Political Economic Perspectives on Health and Disease.\u201d <em>Social Science &amp; Medicine<\/em> 61 (4): 755\u2013765.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Armstrong, C. G., J. E. D. Miller, A. C. McAlvay, P. M. Ritchie, and D. Lepofsky. 2021. \u201cHistorical Indigenous Land-Use Explains Plant Functional Trait Diversity. <em>Ecology and Society<\/em> 26 (2): 6.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Bar-Yosef Mayer, Daniella E., Bernard Vandermeersch, and Ofer Bar-Yosef. 2009. \u201cShells and Ochre in Middle Paleolithic Qafzeh Cave, Israel: Indications for Modern Behavior.\u201d <em>Journal of Human Evolution<\/em> 56 (3): 307\u2013314.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Barbetti, M., and H. Allen. 1972. \u201cPrehistoric Man at Lake Mungo, Australia, by 32,000 Years Bp.\u201d <em>Nature<\/em> 240 (5375): 46\u201348.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Bennett, M. R., D. Bustos, J. S. Pigati, K. B. Springer, T. M. Urban, V. T. Holliday, Sally C. Reynolds, et al. (2021). \u201cEvidence of Humans in North America during the Last Glacial Maximum.\u201d <em>Science<\/em> 373 (6562): 1528\u20131531.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Bowler, J. M., Rhys Jones, Harry Allen, and A. G. Thorne. 1970. \u201cPleistocene Human Remains from Australia: A Living Site and Human Cremation from Lake Mungo, Western New South Wales.\u201d <em>World Archaeology<\/em> 2 (1): 39\u201360.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Brown, Peter. 1999. \u201cThe First Modern East Asians? Another Look at Upper Cave 101, Liujiang and Minatogawa 1.\u201d In <em>Interdisciplinary Perspectives on the Origins of the Japanese<\/em>, edited by K. Omoto, 105\u2013131. Kyoto: International Research Center for Japanese Studies.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Brown, Peter. 2000. \u201cAustralian Pleistocene Variation and the Sex of Lake Mungo 3.\u201d <em>Journal of Human Evolution<\/em> 38 (5): 743\u2013749.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Clarkson, Chris, Zenobia Jacobs, Ben Marwick, Richard Fullagar, Lynley Wallis, Mike Smith, Richard G. Roberts, et al. 2017. \u201cHuman Occupation of Northern Australia by 65,000 Years Ago.\u201d <em>Nature<\/em> 547 (7663): 306\u2013310.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Cohen, Mark Nathan. 1977. <em>The Food Crisis in Prehistory: Overpopulation and the Origins of Agriculture.<\/em> New Haven, CT: Yale University Press.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Cohen, Mark Nathan, and George J. Armelagos, eds. 1984.<em> Paleopathology at the Origins of Agriculture<\/em>. Orlando, FL: Academic Press.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Cohen, Mark Nathan, and Gillian M. M. Crane-Kramer, eds. 2007.<em> Ancient Health: Skeletal Indicators of Agricultural and Economic Intensification<\/em>. Gainesville, FL: University Press of Florida.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Copes-Gerbitz, K., S. Hagerman, and L. Daniels. 2021. \u201cSituating Indigenous Knowledge for Resilience in Fire-Dependent Social-Ecological Systems.\u201d <em>Ecology and Society<\/em> 26(4): 25. https:\/\/www.ecologyandsociety.org\/vol26\/iss4\/art25\/.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Coqueugniot, H\u00e9l\u00e8ne, Olivier Dutour, Baruch Arensburg, Henri Duday, Bernard Vandermeersch, and Anne-Marie Tillier. 2014. \u201cEarliest Cranio-Encephalic Trauma from the Levantine Middle Palaeolithic: 3-D Reappraisal of the Qafzeh 11 Skull, Consequences of Pediatric Brain Damage on Individual Life Condition and Social Care.\u201d <em>PLOS ONE<\/em> 9 (7): e102822.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Crittenden, Alyssa N., and Stephanie L. Schnorr. 2017. \u201cCurrent Views on Hunter\u2010Gatherer Nutrition and the Evolution of the Human Diet.\u201d <em>American Journal of Physical Anthropology<\/em> 162 (S63): 84\u2013109.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">d\u2019Errico, Francesco, Lucinda Backwell, Paola Villa, Ilaria Degano, Jeannette J. Lucejko, Marion K. Bamford, Thomas F. G. Higham, Maria Perla Colombini, and Peter B. Beaumont. 2012. \u201cEarly Evidence of San Material Culture Represented by Organic Artifacts from Border Cave, South Africa.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 109 (33): 13214\u201313219.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">d\u2019Errico, Francesco, Christopher Henshilwood, Marian Vanhaeren, and Karen Van Niekerk. 2005. \u201cNassarius Kraussianus Shell Beads from Blombos Cave: Evidence for Symbolic Behaviour in the Middle Stone Age.\u201d <em>Journal of Human Evolution<\/em> 48 (1): 3\u201324.<\/p>\r\n<p class=\"import-Normal\">Dannemann, Michael, and Fernando Racimo. 2018. \u201cSomething Old, Something Borrowed: Admixture and Adaptation in Human Evolution.\u201d <em>Current Opinion in Genetics &amp; Development<\/em> 53: 1\u20138.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Day, M. H. 1969. \u201cOmo Human Skeletal Remains.\u201d <em>Nature<\/em> 222: 1135\u20131138.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Dillehay, Tom D., Carlos Ocampo, Jos\u00e9 Saavedra, Andre Oliveira Sawakuchi, Rodrigo M. Vega, Mario Pino, Michael B. Collins, et al. 2015. \u201cNew Archaeological Evidence for an Early Human Presence at Monte Verde, Chile.\u201d <em>PLOS ONE<\/em> 10 (11): e0141923. doi:10.1371\/journal.pone.0141923.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Dow, Gregory K., Clyde G. Reed, and Nancy Olewiler. 2009. \u201cClimate Reversals and the Transition to Agriculture.\u201d <em>Journal of Economic Growth<\/em> 14 (1): 27\u201353.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Durband, Arthur C. 2014. \u201cBrief Communication: Artificial Cranial Modification in Kow Swamp and Cohuna.\u201d <em>American Journal of Physical Anthropology<\/em> 155 (1): 173\u2013178.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ember, Carol R. N.d. \u201cHunter-Gatherers.\u201d <em>Explaining Human Culture. Human Relations Area Files<\/em>. Accessed March 4, 2023. <a class=\"rId133\" href=\"https:\/\/hraf.yale.edu\/ehc\/summaries\/hunter-gatherers\">https:\/\/hraf.yale.edu\/ehc\/summaries\/hunter-gatherers<\/a>.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Erlandson, Jon M., Todd J. Braje, Kristina M. Gill, and Michael H. Graham. 2015. \u201cEcology of the Kelp Highway: Did Marine Resources Facilitate Human Dispersal from Northeast Asia to the Americas?\u201d <em>The Journal of Island and Coastal Archaeology<\/em> 10 (3): 392\u2013411.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Fladmark, K. R. 1979. \u201cRoutes: Alternate Migration Corridors for Early Man in North America.\u201d <em>American Antiquity<\/em> 44 (1): 55\u201369.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Fletcher, M. S., T. Hall, and A. N. Alexandra. 2021. \u201cThe Loss of an Indigenous Constructed Landscape Following British Invasion of Australia: An Insight into the Deep Human Imprint on the Australian Landscape.\u201d <em>Ambio<\/em> 50(1): 138\u2013149.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Fu, Qiaomei, Mateja Hajdinjak, Oana Teodora Moldovan, Silviu Constantin, Swapan Mallick, Pontus Skoglund, Nick Patterson, et al. 2015. \u201cAn Early Modern Human from Romania with a Recent Neanderthal Ancestor.\u201d <em>Nature<\/em> 524 (7564): 216\u2013219.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Fuller, Dorian Q. 2010. \u201cAn Emerging Paradigm Shift in the Origins of Agriculture.\u201d <em>General Anthropology<\/em> 17 (2): 1, 8\u201311.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Gammage, B. 2008. \u201cPlain Facts: Tasmania under Aboriginal Management.\u201d <em>Landscape Research<\/em> 33 (2): 241\u2013254.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Germonpr\u00e9, Mietje, Martina L\u00e1zni\u010dkov\u00e1-Galetov\u00e1, and Mikhail V. Sablin. 2012. \u201cPalaeolithic Dog Skulls at the Gravettian P\u0159edmost\u00ed Site, the Czech Republic.\u201d <em>Journal of Archaeological Science<\/em> 39 (1): 184\u2013202.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Gr\u00f6ning, Flora, Jia Liu, Michael J. Fagan, and Paul O\u2019Higgins. 2011. \u201cWhy Do Humans Have Chins? Testing the Mechanical Significance of Modern Human Symphyseal Morphology with Finite Element Analysis.\u201d <em>American Journal of Physical Anthropology<\/em> 144 (4): 593\u2013606.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Harvati, Katerina. 2009. \u201cInto Eurasia: A Geometric Morphometric Reassessment of the Upper Cave (Zhoukoudian) Specimens.\u201d <em>Journal of Human Evolution<\/em> 57 (6): 751\u2013762.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Headland, Thomas N., Lawrence A. Reid, M. G. Bicchieri, Charles A. Bishop, Robert Blust, Nicholas E. Flanders, Peter M. Gardner, Karl L. Hutterer, Arkadiusz Marciniak, and Robert F. Schroeder. 1989. \u201cHunter-Gatherers and Their Neighbors from Prehistory to the Present.\u201d <em>Current Anthropology<\/em> 30 (1): 43\u201366.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Henshilwood, Christopher S., Francesco d\u2019Errico, Karen L. van Niekerk, Yvan Coquinot, Zenobia Jacobs, Stein-Erik Lauritzen, Michel Menu, and Renata Garc\u00eda-Moreno. 2011. \u201cA 100,000-Year-Old Ochre-Processing Workshop at Blombos Cave, South Africa.\u201d <em>Science<\/em> 334 (6053): 219\u2013222.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Hershkovitz, Israel, Gerhard W. Weber, Rolf Quam, Mathieu Duval, Rainer Gr\u00fcn, Leslie Kinsley, Avner Ayalon, et al. 2018. \u201cThe Earliest Modern Humans Outside Africa.\u201d <em>Science<\/em> 359 (6374): 456\u2013459.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Hublin, Jean-Jacques, Abdelouahed Ben-Ncer, Shara E. Bailey, Sarah E. Freidline, Simon Neubauer, Matthew M. Skinner, Inga Bergmann, et al. 2017. \u201cNew Fossils from Jebel Irhoud, Morocco, and the Pan-African Origin of <em>Homo sapiens<\/em>.\u201d <em>Nature<\/em> 546 (7657): 289\u2013292.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Lepofsky, D., N. F. Smith, N. Cardinal, J. Harper, M. Morris, M., Gitla (Elroy White), Randy Bouchard, et al. 2015. \u201cAncient Shellfish Mariculture on the Northwest Coast of North America.\u201d <em>American Antiquity<\/em> 80 (2): 236\u2013259.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Lieberman, Daniel E. 2015. \u201cHuman Locomotion and Heat Loss: An Evolutionary Perspective.\u201d <em>Comprehensive Physiology<\/em> 5 (1): 99\u2013117.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Lieberman, Daniel E., Brandeis M. McBratney, and Gail Krovitz. 2002. \u201cThe Evolution and Development of Cranial Form in <em>Homo sapiens<\/em>.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 99 (3): 1134\u20131139.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Lieberman, Daniel E., Osbjorn M. Pearson, and Kenneth M. Mowbray. 2000. \u201cBasicranial Influence on Overall Cranial Shape.\u201d <em>Journal of Human Evolution<\/em> 38 (2): 291\u2013315.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Liu, Wu, Mar\u00eda Martin\u00f3n-Torres, Yan-jun Cai, Song Xing, Hao-wen Tong, Shu-wen Pei, Mark Jan Sier, Xiao-hong Wu, R. Lawrence Edwards, and Hai Cheng. 2015. \u201cThe Earliest Unequivocally Modern Humans in Southern China.\u201d <em>Nature<\/em> 526 (7575): 696-699.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Lucas, Peter W. 2007. \u201cThe Evolution of the Hominin Diet from a Dental Functional Perspective.\u201d In <em>Evolution of the Human Diet: The Known, the Unknown, and the Unknowable<\/em>, edited by Peter S. Ungar, 31\u201338 Oxford, UK: Oxford University Press.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">McCarthy, Robert C., and Lynn Lucas. 2014. \u201cA Morphometric Reassessment of Bou-Vp-16\/1 from Herto, Ethiopia.\u201d <em>Journal of Human Evolution<\/em> 74: 114\u2013117.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">McDougall, Ian, Francis H. Brown, and John G. Fleagle. 2005. \u201cStratigraphic Placement and Age of Modern Humans from Kibish, Ethiopia.\u201d <em>Nature<\/em> 433 (7027): 733\u2013736.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">McNiven, I. J., J. Crouch, T. Richards, N. Dolby, and G. Jacobsen. 2012. \u201cDating Aboriginal Stone-Walled Fishtraps at Lake Condah, Southeast Australia.\u201d <em>Journal of Archaeological Science<\/em> 39 (2): 268\u2013286.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">McNiven, I., J. Crouch, T. Richards, K. Sniderman, N. Dolby, and G. Mirring. 2015. \u201cPhased Redevelopment of an Ancient Gunditjmara Fish Trap over the Past 800 Years: Muldoons Trap Complex, Lake Condah, Southwestern Victoria.\u201d <em>Australian Archaeology<\/em> 81 (1): 44\u201358.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Michel, V\u00e9ronique, H\u00e9l\u00e8ne Valladas, Guanjun Shen, Wei Wang, Jian-xin Zhao, Chuan-Chou Shen, Patricia Valensi, and Christopher J. Bae. 2016. \u201cThe Earliest Modern <em>Homo sapiens<\/em> in China?\u201d <em>Journal of Human Evolution<\/em> 101: 101\u2013104.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Miller, D. Shane, Vance T. Holliday, and Jordon Bright. 2013. \u201cClovis across the Continent.\u201d In <em>Paleoamerican Odyssey<\/em>, edited by Kelly E. Graf, Caroline V. Ketron, and Michael R. Waters, 207\u2013220. College Station: Texas A&amp;M University Press.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Neubauer, Simon, Jean-Jacques Hublin, and Philipp Gunz. 2018. \u201cThe Evolution of Modern Human Brain Shape.\u201d <em>Science Advances<\/em> 4 (1): eaao5961. https:\/\/doi.org\/10.1126\/sciadv.aao5961.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Pearson, Osbjorn M. 2000. \u201cPostcranial Remains and the Origin of Modern Humans.\u201d <em>Evolutionary Anthropology<\/em> 9: 229\u2013247.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Pearson, Osbjorn M. 2008. \u201cStatistical and Biological Definitions of \u2018Anatomically Modern\u2019 Humans: Suggestions for a Unified Approach to Modern Morphology.\u201d <em>Evolutionary Anthropology: Issues, News, and Reviews<\/em> 17 (1): 38\u201348.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Pietschnig, Jakob, Lars Penke, Jelte M. Wicherts, Michael Zeiler, and Martin Voracek. 2015. \u201cMeta-Analysis of Associations between Human Brain Volume and Intelligence Differences: How Strong Are They and What Do They Mean?\u201d <em>Neuroscience &amp; Biobehavioral Reviews<\/em> 57: 411\u2013432.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Posth, Cosimo, Nathan Nakatsuka, Iosif Lazaridis, Pontus Skoglund, Swapan Mallick, Thiseas C. Lamnidis, Nadin Rohland, et al. 2018. \u201cReconstructing the Deep Population History of Central and South America.\u201d <em>Cell<\/em> 175 (5): 1185\u20131197.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Potter, Ben A., James F. Baichtal, Alwynne B. Beaudoin, Lars Fehren-Schmitz, C. Vance Haynes, Vance T. Holliday, Charles E. Holmes, et al. 2018. \u201cCurrent Evidence Allows Multiple Models for the Peopling of the Americas.\u201d <em>Science Advances<\/em> 4 (8): eaat5473. <a class=\"rId134\" href=\"https:\/\/doi.org\/10.1126\/sciadv.aat5473\">https:\/\/doi.org\/10.1126\/sciadv.aat5473<\/a>.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Reich, David, Richard E. Green, Martin Kircher, Johannes Krause, Nick Patterson, Eric Y. Durand, Bence Viola, et al. 2010. \u201cGenetic History of an Archaic Hominin Group from Denisova Cave in Siberia.\u201d <em>Nature<\/em> 468 (7327): 1053\u20131060.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Reich, David, Nick Patterson, Martin Kircher, Frederick Delfin, Madhusudan R. Nandineni, Irina Pugach, Albert Min-Shan Ko, et al. 2011. \u201cDenisova Admixture and the First Modern Human Dispersals into Southeast Asia and Oceania.\u201d <em>American Journal of Human Genetics<\/em> 89 (4): 516\u2013528.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Richter, Daniel, Rainer Gr\u00fcn, Renaud Joannes-Boyau, Teresa E. Steele, Fethi Amani, Mathieu Ru\u00e9, Paul Fernandes, et al. 2017. \u201cThe Age of the Hominin Fossils from Jebel Irhoud, Morocco, and the Origins of the Middle Stone Age.\u201d <em>Nature<\/em> 546 (7657): 293\u2013296.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Roberts, Patrick, and Brian A. Stewart. 2018. \u201cDefining the \u2018Generalist-Specialist\u2019 Niche for Pleistocene <em>Homo sapiens<\/em>.\u201d <em>Nature Human Behaviour<\/em> 2: 542\u2013550.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Rougier, Helene, \u015etefan Milota, Ricardo Rodrigo, Mircea Gherase, Lauren\u0163iu Sarcin\u01ce, Oana Moldovan, Jo\u00e3o Zilh\u00e3o, et al. 2007. \u201cPe\u015ftera Cu Oase 2 and the Cranial Morphology of Early Modern Europeans.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 104 (4): 1165\u20131170.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Sahlins, Marshall. 1968. \u201cNotes on the Original Affluent Society.\u201d In <em>Man the Hunter<\/em>, edited by R. B. Lee and I. DeVore, 85\u201389. New York: Aldine Publishing Company.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Sawyer, G. J., and Blaine Maley. 2005. \u201cNeanderthal Reconstructed.\u201d <em>The Anatomical Record (Part B: New Anat.)<\/em> 283 (1): 23\u201331.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scerri, Eleanor M. L., Mark G. Thomas, Andrea Manica, Philipp Gunz, Jay T. Stock, Chris Stringer, Matt Grove, et al. 2018. \u201cDid Our Species Evolve in Subdivided Populations Across Africa, and Why Does It Matter?\u201d <em>Trends in Ecology &amp; Evolution<\/em> 33 (8): 582\u2013594.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Shea, John J. 2011. \u201cRefuting a Myth about Human Origins.\u201d <em>American Scientist<\/em> 99 (2): 128\u2013135.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Shea, John J., and Ofer Bar-Yosef. 2005. \u201cWho Were the Skhul\/Qafzeh People? An Archaeological Perspective on Eurasia\u2019s Oldest Modern Humans.\u201d <em>Journal of the Israel Prehistoric Societ<\/em><em>y<\/em> 35: 451\u2013468.<\/p>\r\n<p class=\"import-Normal\">Slatkin, Montgomery, and Fernando Racimo. 2016. \u201cAncient DNA and Human History.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 113 (23): 6380\u20136387.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Smith, Fred H., James C. M. Ahern, Ivor Jankovi\u0107, and Ivor Karavani\u0107. 2017. \u201cThe Assimilation Model of Modern Human Origins in Light of Current Genetic and Genomic Knowledge.\u201d <em>Quaternary International<\/em> 450: 126\u2013136.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Smith, Michael. 2009. \u201cV. Gordon Childe and the Urban Revolution: A Historical Perspective on a Revolution in Urban Studies.\u201d <em>Town Planning Review<\/em> 80 (1): 3\u201329.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Stock, Jay T. 2008. \u201cAre Humans Still Evolving?\u201d <em>EMBO Reports<\/em> 9 (Suppl 1): S51\u2013S54.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Swisher, Mark E., Dennis L. Jenkins, Lionel E. Jackson Jr., and Fred M. Phillips. 2013. \u201cA Reassessment of the Role of the Canadian Ice-Free Corridor in Light of New Geological Evidence.\u201d Poster Symposium 5B: Geology, Geochronology and Paleoenvironments of the First Americans at the Paleoamerican Odyssey Conference, Santa Fe, New Mexico, October 16\u201319.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Thorne, A. G., and P. G. Macumber. 1972. \u201cDiscoveries of Late Pleistocene Man at Kow Swamp, Australia.\u201d <em>Nature<\/em> 238 (5363): 316\u2013319.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Trinkaus, Erik, \u015etefan Milota, Ricardo Rodrigo, Gherase Mircea, and Oana Moldovan. 2003a. \u201cEarly Modern Human Cranial Remains from the Pe\u015ftera Cu Oase, Romania.\u201d <em>Journal of Human Evolution<\/em> 45 (3): 245\u2013253.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Trinkaus, Erik, Oana Moldovan, Adrian B\u00eelg\u0103r, Lauren\u0163iu Sarcina, Sheela Athreya, Shara E Bailey, Ricardo Rodrigo, Gherase Mircea, Thomas Higham, and Christopher Bronk Ramsey. 2003b. \u201cAn Early Modern Human from the Pe\u015ftera Cu Oase, Romania.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 100 (20): 11231\u201311236.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Velem\u00ednsk\u00e1, J., J. Br\u016fzek, P. Velem\u00ednsk\u00fd, L. Bigoni, A. Sefc\u00e1kov\u00e1, and S. Katina. 2008. \u201cVariability of the Upper-Palaeolithic Skulls from Predmost\u00ed Near Prerov (Czech Republic): Craniometric Comparison with Recent Human Standards.\u201d <em>Homo<\/em> 59 (1): 1\u201326.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Vidal, C\u00e9line M., Christine S. Lane, Asfawossen Asrat, Dan N. Barfod, Darren F. Mark, Emma L. Tomlinson, Ambdemichael Zafu Tadesse, et al. (2022). \u201cAge of the Oldest Known <em>Homo sapiens<\/em> from Eastern Africa. <em>Nature<\/em> 601 (7894): 579\u2013583.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Villa, Paola, Sylvain Soriano, Tsenka Tsanova, Ilaria Degano, Thomas F. G. Higham, Francesco d\u2019Errico, Lucinda Backwell, Jeannette J. Lucejko, Maria Perla Colombini, and Peter B. Beaumont. 2012. \u201cBorder Cave and the Beginning of the Later Stone Age in South Africa.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 109 (33): 13208\u201313213.<\/p>\r\n<p class=\"import-Normal\">Wall, Jeffrey D., and Deborah Yoshihara Caldeira Brandt. 2016. \u201cArchaic Admixture in Human History.\u201d <em>Current Opinion in Genetics &amp; Development<\/em> 41: 93\u201397.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">White, Tim D., Berhane Asfaw, David DeGusta, Henry Gilbert, Gary D. Richards, Gen Suwa, and F. Clark Howell. 2003. \u201cPleistocene <em>Homo sapiens<\/em> from Middle Awash, Ethiopia.\u201d <em>Nature<\/em> 423 (6941): 742\u2013747.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Woo, Ju-Kang. 1959. \u201cHuman Fossils Found in Liukiang, Kwangsi, China.\u201d <em>Vertebrata PalAsiatica<\/em> 3 (3): 109\u2013118.<\/p>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Wu, XiuJie, Wu Liu, Wei Dong, JieMin Que, and YanFang Wang. 2008. \u201cThe Brain Morphology of Homo Liujiang Cranium Fossil by Three-Dimensional Computed Tomography.\u201d <em>Chinese Science Bulletin<\/em> 53 (16): 2513\u20132519.<\/p>\r\n\r\n<h2 class=\"import-Normal\">Acknowledgments<\/h2>\r\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">I could not have undertaken this project without the help of many who got me to where I am today. I extend sincere thank yous to the many colleagues and former students who have inspired me to keep learning and talking about anthropology. Thank you also to all who are involved in this textbook project. The anonymous reviewers truly sparked improvements to the chapter. Lastly, the staff of Starbucks #5772 also contributed immensely to this text.<\/p>\r\n\r\n<\/div>","rendered":"<div class=\"__UNKNOWN__\">\n<p>Keith Chan, Ph.D., Grossmont-Cuyamaca Community College District and MiraCosta College<\/p>\n<p><em>This chapter is a revision from &#8220;<\/em><a class=\"rId7\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-14\/\"><em>Chapter 12: Modern Homo sapiens<\/em><\/a><em>\u201d by Keith Chan. In <\/em><a class=\"rId8\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, first edition<\/em><\/a><em>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under <\/em><a class=\"rId9\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\"><em>CC BY-NC 4.0<\/em><\/a><em>. <\/em><\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\"><span style=\"color: #ffffff\">Learning Objectives<\/span><\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li>Identify the skeletal and behavioral traits that represent modern <em>Homo sapiens.<\/em><\/li>\n<li>Critically evaluate different types of evidence for the origin of our species in Africa and our expansion around the world.<\/li>\n<li>Understand how the human lifestyle changed when people transitioned from foraging to agriculture.<\/li>\n<li>Hypothesize how human evolutionary trends may continue into the future.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p class=\"import-Normal\">The walls of a pink limestone cave in the hillside of Jebel Irhoud jutted out of the otherwise barren landscape of the Moroccan desert (Figure 12.1). Miners had excavated the cave in the 1960s, revealing some fossils. In 2007, a re-excavation of the site became a momentous occasion for science. A fossil cranium unearthed by a team of researchers was barely visible to the untrained eye. Just the fossil\u2019s robust brows were peering out of the rock. This research team from the Max Planck Institute for Evolutionary Anthropology was the latest to explore the ancient human presence in this part of North Africa after a find by miners in 1960. Excavating near the first discovery, the researchers wanted to learn more about how <em>Homo sapiens<\/em> lived far from East Africa, where we thought our species originated.<\/p>\n<figure style=\"width: 2500px\" class=\"wp-caption alignnone\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/06\/image10-1.jpg\" alt=\"Rocky hillside with exposed layers. People are visible at the base.\" width=\"2500\" height=\"987\" \/><figcaption class=\"wp-caption-text\">Figure 12.1: The excavation of an exposed cave at Jebel Irhoud, Morocco, where hominin fossils were found in the 1960s and in 2007. Dating showed that they could represent the earliest-known modern Homo sapiens. Credit: <a href=\"https:\/\/www.eva.mpg.de\/homo-sapiens\/presskit.html\">View looking south of the Jebel Irhoud (Morocco) site<\/a> by Shannon McPherron, <a href=\"https:\/\/www.eva.mpg.de\/index.html\">MPI EVA Leipzig<\/a>, is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/\">CC BY-SA 2.0 License<\/a>.<\/figcaption><\/figure>\n<p>The scientists were surprised when they analyzed the cranium, named Irhoud 10, and other fossils. Statistical comparisons with other human crania concluded that the Irhoud face shapes were typical of recent modern humans while the braincases matched ancient modern humans. Based on the findings of other scientists, the team expected these modern <em>Homo sapiens<\/em> fossils to be around 200,000 years old. Instead, dating revealed that the cranium had been buried for around 315,000 years.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Together, the modern-looking facial dimensions and the older date reshaped the interpretation of our species: modern <em>Homo sapiens<\/em>. Some key evolutionary changes from the archaic <em>Homo sapiens<\/em> (described in Chapter 11) to our species today happened 100,000 years earlier than we had thought and across the vast African continent rather than concentrated in its eastern region.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">This revelation in the study of modern <em>Homo sapiens<\/em> is just one of the latest in this continually advancing area of biological anthropology. Researchers today are still discovering amazing fossils and ingenious ways to collect data and test hypotheses about our past. Through the collective work of many scientists, we are building an overall theory of modern human origins. <span style=\"background-color: #ff99cc\">In this chapter, we will first cover the skeletal changes from archaic <em>Homo sapiens<\/em> to modern <em>Homo sapiens<\/em>. Next, we will track how modern <em>Homo sapiens<\/em> expanded around the world. Lastly, we will cover the development of agriculture and how it changed human culture.<\/span><\/p>\n<h2 class=\"import-Normal\">Defining Modernity<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">What defines modern <em>Homo sapiens<\/em> when compared to archaic <em>Homo sapiens<\/em>? Modern humans, like you and me, have a set of derived traits that are not seen in archaic humans or any other hominin. As with other transitions in hominin evolution, such as increasing brain size and bipedal ability, modern traits do not appear fully formed or all at once. In other words, the first modern <em>Homo sapiens<\/em> was not just born one day from archaic parents. The traits common to modern <em>Homo sapiens<\/em> appeared in a <strong>mosaic<\/strong> manner: gradually and out of sync with one another. There are two areas to consider when tracking the complex evolution of modern human traits. One is the physical change in the skeleton. The other is behavior inferred from the size and shape of the cranium and material culture evidence.<\/p>\n<h3 class=\"import-Normal\"><strong>Skeletal Traits<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The skeleton of modern <em>Homo sapiens<\/em> is less robust than that of archaic <em>Homo sapiens<\/em>. In other words, the modern skeleton is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1406\">gracile<\/a><\/strong>, meaning that the structures are thinner and smoother. Differences related to gracility in the cranium are seen in the braincase, the face, and the mandible. There are also broad differences in the rest of the skeleton.<\/p>\n<h4 class=\"import-Normal\"><em>Cranial Traits<\/em><\/h4>\n<figure style=\"width: 445px\" class=\"wp-caption alignleft\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image29-2.png\" alt=\"A rounded skull facing a robust skull with sloping forehead.\" width=\"445\" height=\"221\" \/><figcaption class=\"wp-caption-text\">Figure 12.2: Comparison between modern (left) and archaic (right) Homo sapiens skulls. Note the overall gracility of the modern skull, as well as the globular braincase. Credit: <a class=\"rId15\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-14\/\">Modern human and Neanderthal<\/a> original to <a class=\"rId16\" href=\"https:\/\/explorations.americananthro.org\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a class=\"rId17\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Several elements of the braincase differ between modern and archaic <em>Homo sapiens<\/em>. Overall, the shape is much rounder, or more <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1758\"><strong>globular<\/strong>,<\/a> on a modern skull (Lieberman, McBratney, and Krovitz 2002; Neubauer, Hublin, and Gunz 2018; Pearson 2008; Figure 12.2). You can feel the globularity of your own modern human skull. Feel the height of your forehead with the palm of your hand. Viewed from the side, the tall vertical forehead of a modern <em>Homo sapiens<\/em> stands out when compared to the sloping archaic version. This is because the frontal lobe of the modern human brain is larger than the one in archaic humans, and the skull has to accommodate the expansion. The vertical forehead reduces a trait that is common to all other hominins: the brow ridge or <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1759\">supraorbital torus<\/a><\/strong>. The parietal lobes of the brain and the matching parietal bones on either side of the skull both bulge outward more in modern humans. At the back of the skull, the archaic occipital bun is no longer present. Instead, the occipital region of the modern human cranium has a derived tall and smooth curve, again reflecting the globular brain inside.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The trend of shrinking face size across hominins reaches its extreme with our species as well. The facial bones of a modern <em>Homo sapiens<\/em> are extremely gracile compared to all other hominins (Lieberman, McBratney, and Krovitz 2002). Continuing a trend in hominin evolution, technological innovations kept reducing the importance of teeth in reproductive success (Lucas 2007). As natural selection favored smaller and smaller teeth, the surrounding bone holding these teeth also shrank.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Related to smaller teeth, the mandible is also gracile in modern humans when compared to archaic humans and other hominins. Interestingly, our mandibles have pulled back so far from the prognathism of earlier hominins that we gained an extra structure at the most anterior point, called the <strong>mental eminence<\/strong>. You know this structure as the chin. At the skeletal level, it resembles an upside-down \u201cT\u201d at the centerline of the mandible (Pearson 2008). Looking back at archaic humans, you will see that they all lack a chin. Instead, their mandibles curve straight back without a forward point. What is the chin for and how did it develop? Flora Gr\u00f6ning and colleagues (2011) found evidence of the chin\u2019s importance by simulating physical forces on computer models of different mandible shapes. Their results showed that the chin acts as structural support to withstand strain on the otherwise gracile mandible.<\/p>\n<h4 class=\"import-Normal\"><em>Postcranial Gracility<\/em><\/h4>\n<figure style=\"width: 368px\" class=\"wp-caption alignright\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image16-5.png\" alt=\"Two complete skeletons. The left is taller with a thinner frame.\" width=\"368\" height=\"575\" \/><figcaption class=\"wp-caption-text\">Figure 12.3: Anterior views of modern (left) and archaic (right) Homo sapiens skeletons. The modern human has an overall gracile appearance at this scale as well. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-14\/\">Modern and archaic Homo sapiens skeletons (Figure 12.3)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p>The rest of the modern human skeleton is also more gracile than its archaic counterpart. The differences are clear when comparing a modern <em>Homo sapiens<\/em> with a cold-adapted Neanderthal (Sawyer and Maley 2005), but the trends are still present when comparing modern and archaic humans within Africa (Pearson 2000). Overall, a modern <em>Homo sapiens<\/em> postcranial skeleton has thinner cortical bone, smoother features, and more slender shapes when compared to archaic <em>Homo sapiens<\/em> (Figure 12.3). Comparing whole skeletons, modern humans have longer limb proportions relative to the length and width of the torso, giving us lankier outlines.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Why is our skeleton so gracile compared to those of other hominins? Natural selection can drive the gracilization of skeletons in several ways (Lieberman 2015). <span style=\"background-color: #ffff00\">A slender frame is adapted for the efficient long-distance running ability that started with <em>Homo erectus<\/em>. Furthermore, slenderness is a genetic adaptation for cooling an active body in hotter climates, which aligns with the ample evidence that Africa was the home continent of our species.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong>Behavioral Modernity<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Aside from physical differences in the skeleton, researchers have also uncovered evidence of behavioral changes associated with increased cultural complexity from archaic to modern humans. How did cultural complexity develop? Two investigations into this question are archaeology and the analysis of reconstructed brains.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Archaeology tells us much about the behavioral complexity of past humans by interpreting the significance of material culture. In terms of advanced culture, items created with an artistic flair, or as decoration, speak of abstract thought processes (Figure 12.4). The demonstration of difficult artistic techniques and technological complexity hints at social learning and cooperation as well. According to paleoanthropologist John Shea (2011), one way to track the complexity of past behavior through artifacts is by measuring the variety of tools found together. The more types of tools constructed with different techniques and for different purposes, the more modern the behavior. Researchers are still working on an archaeological way to measure cultural complexity that is useful across time and place.<\/p>\n<figure style=\"width: 221px\" class=\"wp-caption alignleft\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image15-1-1.jpg\" alt=\"A brown standing statue of a human figure with cat\u2019s head.\" width=\"221\" height=\"392\" \/><figcaption class=\"wp-caption-text\">Figure 12.4: Carved ivory figure called \u201cthe Lion-Man of the Hohlenstein-Stadel.\u201d It dates to the Aurignacian culture, between 35 and 40 kya. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Loewenmensch1.jpg\">Loewenmensch1<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Dagmar_Hollmann\">Dagmar Hollmann<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The interpretation of brain anatomy is another promising approach to studying the evolution of human behavior. When looking at investigations on this topic in modern <em>Homo sapiens<\/em> brains, researchers found a weak association between brain size and test-measured intelligence (Pietschnig et al. 2015). Additionally, they found no association between intelligence and biological sex. These findings mean that there are more significant factors that affect tested intelligence than just brain size. Since the sheer size of the brain is not useful for weighing intelligence within a species, paleoanthropologists are instead investigating the differences in certain brain structures. The differences in organization between modern <em>Homo sapiens<\/em> brains and archaic <em>Homo sapiens<\/em> brains may reflect different cognitive priorities that account for modern human culture. As with the archaeological approach, new discoveries will refine what we know about the human brain and apply that knowledge to studying the distant past.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Taken together, the cognitive abilities in modern humans may have translated into an adept use of tools to enhance survival. Researchers Patrick Roberts and Brian A. Stewart (2018) call this concept the <strong>generalist-specialist niche<\/strong>: our species is an expert at living in a wide array of environments, with populations culturally specializing in their own particular surroundings. The next section tracks how far around the world these skeletal and behavioral traits have taken us.<\/p>\n<h2 class=\"import-Normal\">First Africa, Then the World<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">What enabled modern <em>Homo sapiens<\/em> to expand its range further in 300,000 years than <em>Homo erectus<\/em> did in 1.5 million years? <span style=\"background-color: #ffff00\">The key is the set of derived biological traits from the last section. The gracile frame and neurological anatomy allowed modern humans to survive and even flourish in the vastly different environments they encountered.<\/span> Based on multiple types of evidence, the source of all of these modern humans was Africa. Instead of originating from just one location, evidence shows that modern Homo sapiens evolution occurred in a complex gene flow network across Africa, a concept called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1760\">African multiregionalism<\/a><\/strong> (Scerri et al. 2018).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">This section traces the origin of modern <em>Homo sapiens<\/em> and the massive expansion of our species across all of the continents (except Antarctica) by 12,000 years ago. While modern <em>Homo sapiens<\/em> first shared geography with archaic humans, modern humans eventually spread into lands where no human had gone before. Figure 12.5 shows the broad routes that our species took expanding around the world. I encourage you to make your own timeline with the dates in this part to see the overall trends.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image13-6.png\" alt=\"315 to 195 KYA. Northern to eastern coasts of Africa are shaded.\" width=\"554\" height=\"428\" \/><\/p>\n<p class=\"import-Normal\"><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image17-5.png\" alt=\"195-100 KYA. Africa, southern Europe and Asia are shaded\" width=\"554\" height=\"428\" \/><\/p>\n<p class=\"import-Normal\"><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image27-3.png\" alt=\"99 to 30 KYA. Africa, Indonesia, Australia, and southern portions of Europe and Asia are shaded.\" width=\"554\" height=\"428\" \/><\/p>\n<figure style=\"width: 554px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image30-2.png\" alt=\"29 to 9 KYA. Shading covers most land except Antarctica, Greenland, and some islands.\" width=\"554\" height=\"428\" \/><figcaption class=\"wp-caption-text\">Figure 12.5a-d: Four maps depicting the estimated range of modern Homo sapiens through time. The shaded area is based on geographical connections across known sites. Note the growth in the area starting in Africa and the oftentimes-coastal routes that populations followed. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-14\/\">Four maps depicting the estimated range of modern Homo sapiens through time<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Elyssa Ebding at <a href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\">GeoPlace, California State University, Chico<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Modern <\/strong><strong><em>Homo sapiens<\/em><\/strong><strong> Biology and Culture in Africa<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">We start with the ample fossil evidence supporting the theory that modern humans originated in Africa during the Middle Pleistocene, having evolved from African archaic <em>Homo sapiens<\/em>. The earliest dated fossils considered to be modern actually have a mosaic of archaic and modern traits, showing the complex changes from one type to the other. Experts have various names for these transitional fossils, such as <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1763\"><strong>Early Modern <\/strong><strong><em>Homo sapiens\u00a0 <\/em><\/strong><\/a><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1764\"> or Early Anatomically Modern Humans<\/a><\/strong>. However they are labeled, the presence of some modern traits means that they illustrate the origin of the modern type. Three particularly informative sites with fossils of the earliest modern <em>Homo sapiens<\/em> are Jebel Irhoud, Omo, and Herto.<\/p>\n<figure style=\"width: 281px\" class=\"wp-caption alignleft\"><img loading=\"lazy\" decoding=\"async\" class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image9-1-1.jpg\" alt=\"3D image of a human cranium with pronounced brow ridges.\" width=\"281\" height=\"282\" \/><figcaption class=\"wp-caption-text\">Figure 12.6: Composite rendering of the Jebel Irhoud hominin based on micro-CT scans of multiple fossils from the site. The facial structure is within the modern human range, while the braincase is between the archaic and modern shapes. Credit: <a href=\"https:\/\/www.eva.mpg.de\/homo-sapiens\/presskit.html\">A composite reconstruction of the earliest known Homo sapiens fossils from Jebel Irhoud (Morocco) based on micro computed tomographic scans<\/a> by Philipp Gunz, <a href=\"https:\/\/www.eva.mpg.de\/index.html\">MPI EVA Leipzig<\/a>, is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/\">CC BY-SA 2.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Recall from the start of the chapter that the most recent finds at Jebel Irhoud are now the oldest dated fossils that exhibit some facial traits of modern <em>Homo sapiens<\/em>. Besides Irhoud 10, the cranium that was dated to 315,000 years ago (Hublin et al. 2017; Richter et al. 2017), there were other fossils found in the same deposit that we now know are from the same time period. In total there are at least five individuals, representing life stages from childhood to adulthood. These fossils form an image of high variation in skeletal traits. For example, the skull named Irhoud 1 has a primitive brow ridge, while Irhoud 2 and Irhoud 10 do not (Figure 12.6). The braincases are lower than what is seen in the modern humans of today but higher than in archaic <em>Homo sapiens<\/em>. The teeth also have a mix of archaic and modern traits that defy clear categorization into either group.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Research separated by nearly four decades uncovered fossils and artifacts from the Kibish Formation in the Lower Omo Valley in Ethiopia. These Omo Kibish hominins were represented by braincases and fragmented postcranial bones of three individuals found kilometers apart, dating back to around 233,000 years ago (Day 1969; McDougall, Brown, and Fleagle 2005; Vidal et al. 2022). One interesting finding was the variation in braincase size between the two more-complete specimens: while the individual named Omo I had a more globular dome, Omo II had an archaic-style long and low cranium.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Also in Ethiopia, a team led by Tim White (2003) excavated numerous fossils at Herto. There were fossilized crania of two adults and a child, along with fragments of more individuals. The dates ranged between 160,000 and 154,000 years ago. The skeletal traits and stone-tool assemblage were both intermediate between the archaic and modern types. Features reminiscent of modern humans included a tall braincase and thinner zygomatic (cheek) bones than those of archaic humans (Figure 12.7). Still, some archaic traits persisted in the Herto fossils, such as the supraorbital tori. Statistical analysis by other research teams concluded that at least some cranial measurements fit just within the modern human range (McCarthy and Lucas 2014), favoring categorization with our own species.<\/p>\n<figure style=\"width: 373px\" class=\"wp-caption alignright\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image4-3.jpg\" alt=\"Replica cranium showing wide brow ridges and gracile face.\" width=\"373\" height=\"373\" \/><figcaption class=\"wp-caption-text\">Figure 12.7: This model of the Herto cranium showing its mosaic of archaic and modern traits. Credit: <a href=\"https:\/\/boneclones.com\/product\/homo-sapiens-idaltu-bou-vp-16-1-herto-skull-BH-045\/category\/all-fossil-hominids\/fossil-hominids\">Homo sapiens idaltu BOU-VP-16\/1 Herto Cranium<\/a> by <a href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a> is used by permission and available here under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">The timeline of material culture suggests a long period of relying on similar tools before a noticeable diversification of artifacts types. Researchers label the time of stable technology shared with archaic types the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1766\"><strong>Middle Stone Age<\/strong><\/a>, while the subsequent time of diversification in material culture is called the<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1767\"> <strong>Later Stone Age<\/strong><\/a>.<\/p>\n<p class=\"import-Normal\">In the Middle Stone Age, the sites of Jebel Irhoud, Omo, and Herto all bore tools of the same flaked style as archaic assemblages, even though they were separated by almost 150,000 years. The consistency in technology may be evidence that behavioral modernity was not so developed. No clear signs of art dating back this far have been found either. Other hypotheses not related to behavioral modernity could explain these observations. The tool set may have been suitable for thriving in Africa without further innovation. Maybe works of art from that time were made with media that deteriorated or perhaps such art was removed by later humans.<\/p>\n<p class=\"import-Normal\">Evidence of what <em>Homo sapiens<\/em> did in Africa from the end of the Middle Stone Age to the Later Stone Age is concentrated in South African cave sites that reveal the complexity of human behavior at the time. For example, Blombos Cave, located along the present shore of the Cape of Africa facing the Indian Ocean, is notable for having a wide variety of artifacts. The material culture shows that toolmaking and artistry were more complex than previously thought for the Middle Stone Age. In a layer dated to 100,000 years ago, researchers found two intact ochre-processing kits made of abalone shells and grinding stones (Henshilwood et al. 2011). Marine snail shell beads from 75,000 years ago were also excavated (Figure 12.8; d\u2019Errico et al. 2005). Together, the evidence shows that the Middle Stone Age occupation at Blombos Cave incorporated resources from a variety of local environments into their culture, from caves (ochre), open land (animal bones and fat), and the sea (abalone and snail shells). This complexity shows a deep knowledge of the region\u2019s resources and their use\u2014not just for survival but also for symbolic purposes.<\/p>\n<figure style=\"width: 563px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image5-2-1.jpg\" alt=\"Multiple views of shells with holes bored through them.\" width=\"563\" height=\"482\" \/><figcaption class=\"wp-caption-text\">Figure 12.8: Examples of the perforated shell beads found in Blombos Cave, South Africa: (a) view of carved hole seen from the inside; (b) arrows indicate worn surfaces due to repetitive contact with other objects, such as with other beads or a connecting string; (c) traces of ochre; and (d) four shell beads showing a consistent pattern of perforation. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:BBC-shell-beads.jpg\">BBC-shell-beads<\/a> by Chenshilwood (Chris Henshilbood and Francesco d\u2019Errico) at <a href=\"https:\/\/en.wikipedia.org\/wiki\/\">English Wikipedia<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">On the eastern coast of South Africa, Border Cave shows new African cultural developments at the start of the Later Stone Age. Paola Villa and colleagues (2012) identified several changes in technology around 43,000 years ago. Stone-tool production transitioned from a slower process to one that was faster and made many <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1777\">microliths<\/a><\/strong>, small and precise stone tools. Changes in decorations were also found across the Later Stone Age transition. Beads were made from a new resource: fragments of ostrich eggs shaped into circular forms resembling present-day breakfast cereal O\u2019s (d\u2019Errico et al. 2012). These beads show a higher level of altering one\u2019s own surroundings and a move from the natural to the abstract in terms of design.<\/p>\n<h4 class=\"import-Normal\"><span style=\"background-color: #ff99cc\"><em>Summary of Modern <\/em>H. sapiens<em> in Africa<\/em><\/span><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">The combined fossil evidence paints a picture of diversity in geography and traits. Instead of evolving in just East Africa, the Jebel Irhoud find revealed that early modern <em>Homo sapiens<\/em> had a wide range across Middle Pleistocene Africa. Supporting this explanation, fossils have different mosaics of archaic and modern traits in different places and even within the same area. The high level of diversity from just these fossils shows that the modern traits took separate paths toward the set we have today. The connections were convoluted, involving fluctuating gene flow among small groups of regional nomadic foragers across a large continent over a long time.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ff99cc\">African culture experienced a long constant phase called the Middle Stone Age until a faster burst of change produced innovation and new styles. The change was not one moment but rather an escalation in development. Later Stone Age culture introduced elements seen across many regions, including the construction of composite tools and even the use of strung decorations such as beads. These developments appear in the Later Stone Age of other regions, such as Europe. Based on the early date of the African artifacts, Later Stone Age culture may have originated in Africa and passed from person to person and region to region, with people adapting the general technique to their local resources and viewing the meaning in their own way.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong>Expansion into the Middle East and Asia<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">While modern <em>Homo sapiens<\/em> lived across Africa, some members eventually left the continent. These pioneers could have used two connections to the Middle East or West Asia. From North Africa, they could have crossed the Sinai Peninsula and moved north to the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1769\"><strong>Levant<\/strong><\/a>, or eastern Mediterranean. Finds in that region show an early modern human presence. Other finds support the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1770\">Southern Dispersal model<\/a><\/strong>, with a crossing from East Africa to the southern Arabian Peninsula through the Straits of Bab-el-Mandeb. It is tempting to think of one momentous event in which people stepped off Africa and into the Middle East, never to look back. In reality, there were likely multiple waves of movement producing gene flow back and forth across these regions as the overall range pushed east. The expanding modern human population could have thrived by using resources along the southern coast of the Arabian Peninsula to South Asia, with side routes moving north along rivers. The maximum range of the species then grew across Asia.<\/p>\n<h4 class=\"import-Normal\"><em>Modern <\/em>Homo sapiens<em> in the Middle East<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Geographically, the Middle East is the ideal place for the African modern <em>Homo sapiens<\/em> population to inhabit upon expanding out of their home continent. In the Eastern Mediterranean coast of the Levant, there is a wealth of skeletal and material culture linked to modern <em>Homo sapiens<\/em>. Recent discoveries from Saudi Arabia further add to our view of human life just beyond Africa.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The Caves of Mount Carmel in present-day Israel have preserved skeletal remains and artifacts of modern <em>Homo sapiens<\/em>, the first-known group living outside Africa. The skeletal presence at Misliya Cave is represented by just part of the left upper jaw of one individual, but it is notable for being dated to a very early time, between 194,000 and 177,000 years ago (Hershkovitz et al. 2018). Later, from 120,000 to 90,000 years ago, fossils of multiple individuals across life stages were found in the caves of Es-Skhul and Qafzeh (Shea and Bar-Yosef 2005). The skeletons had many modern <em>Homo sapiens<\/em> traits, such as globular crania and more gracile postcranial bones when compared to Neanderthals. Still, there were some archaic traits. For example, the adult male Skhul V also possessed what researchers Daniel Lieberman, Osbjorn Pearson, and Kenneth Mowbray (2000) called marked or clear occipital bunning. Also, compared to later modern humans, the Mount Carmel people were more robust. Skhul V had a particularly impressive brow ridge that was short in height but sharply jutted forward above the eyes (Figure 12.9). The high level of preservation is due to the intentional burial of some of these people. Besides skeletal material, there are signs of artistic or symbolic behavior. For example, the adult male Skhul V had a boar\u2019s jaw on his chest. Similarly, Qafzeh 11, a juvenile with healed cranial trauma, had an impressive deer antler rack placed over his torso (Figure 12.10; Coqueugniot et al. 2014). Perforated seashells colored with <strong>ochre<\/strong>, mineral-based pigment, were also found in Qafzeh (Bar-Yosef Mayer, Vandermeersch, and Bar-Yosef 2009).<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 484px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image6-2-1.jpg\" alt=\"Side view of a skull replica with a globular braincase.\" width=\"484\" height=\"484\" \/><figcaption class=\"wp-caption-text\">Figure 12.9: This Skhul V cranium model shows the sharp browridges. The contour of a marked occipital bun is barely visible from this angle. Credit: <a href=\"https:\/\/boneclones.com\/product\/homo-sapiens-skull-skhul-5-BH-032\">Homo sapiens Skull Skhul 5<\/a> by <a href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a> is used by permission and available here under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p>&nbsp;<\/p>\n<figure style=\"width: 484px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image26-1-1.jpg\" alt=\"Human skeleton in a stony matrix. Ribs are visible below the antlers.\" width=\"484\" height=\"312\" \/><figcaption class=\"wp-caption-text\">Figure 12.10 This cast of the Qafzeh 11 burial shows the antler\u2019s placement over the upper torso. The forearm bones appear to overlap the antler. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Moulage_de_la_s%C3%A9pulture_de_l'individu_%22Qafzeh_11%22_(avec_ramure_de_cervid%C3%A9),_homme_de_N%C3%A9andertal.jpg\">Moulage de la s\u00e9pulture de l&#8217;individu &#8220;Qafzeh 11&#8221; (avec ramure de cervid\u00e9), homme de N\u00e9andertal<\/a> (Collections du Mus\u00e9um national d&#8217;histoire naturelle de Paris, France) by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Eunostos\">Eunostos<\/a> has been modified (cropped and color modified) and is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">One remaining question is, what happened to the modern humans of the Levant after 90,000 years ago? Another site attributed to our species did not appear in the region until 47,000 years ago. Competition with Neanderthals may have accounted for the disappearance of modern human occupation since the Neanderthal presence in the Levant lasted longer than the dates of the early modern <em>Homo sapiens<\/em>. John Shea and Ofer Bar-Yosef (2005) hypothesized that the Mount Carmel modern humans were an initial expansion from Africa that failed. Perhaps they could not succeed due to competition with the Neanderthals who had been there longer and had both cultural and biological adaptations to that environment.<\/p>\n<h4 class=\"import-Normal\"><em>Modern <\/em>Homo sapiens<em> of China<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">A long history of paleoanthropology in China has found ample evidence of modern human presence. Four notable sites are the caves at Fuyan, Liujiang, Tianyuan, and Zhoukoudian. In the distant past, these caves would have been at least seasonal shelters that unintentionally preserved evidence of human presence for modern researchers to discover.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">At Fuyan Cave in Southern China, paleoanthropologists found 47 adult teeth associated with cave formations dated to between 120,000 and 80,000 years ago (Liu et al. 2015). It is currently the oldest-known modern human site in China, though other researchers question the validity of the date range (Michel et al. 2016). The teeth have the small size and gracile features of modern <em>Homo sapiens<\/em> dentition.<\/p>\n<p class=\"import-Normal\">The fossil Liujiang (or Liukiang) hominin (67,000 years ago) has derived traits that classified it as a modern <em>Homo sapiens<\/em>, though primitive archaic traits were also present. In the skull, which was found nearly complete, the Liujiang hominin had a taller forehead than archaic <em>Homo sapiens<\/em> but also had an enlarged occipital region (Figure 12.11; Brown 1999; Wu et al. 2008). Other parts of the skeleton also had a mix of modern and archaic traits: for example, the femur fragments suggested a slender length but with thick bone walls (Woo 1959).<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 486px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image19-1-2.jpg\" alt=\"A human skull with very slight brow ridges and an extremely globular braincase.\" width=\"486\" height=\"323\" \/><figcaption class=\"wp-caption-text\">Figure 12.11: The Liujiang cranium shows the tall forehead and overall gracile appearance typical of modern Homo sapiens. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Liujiang_cave_skull-a._Homo_Sapiens_68,000_Years_Old.jpg\">Liujiang cave skull-a. Homo Sapiens 68,000 Years Old<\/a> (Taken at the David H. Koch Hall of Human Origins, <a href=\"https:\/\/naturalhistory.si.edu\/visit\">Smithsonian Natural History Museum<\/a>) by <a href=\"https:\/\/www.flickr.com\/people\/14405058@N08\">Ryan Somma<\/a> has been modified (color modified) and is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/legalcode\">CC BY-SA 2.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Another Chinese site to describe here is the one that has been studied the longest. In the Zhoukoudian Cave system (Figure 12.12), where <em>Homo erectus<\/em> and archaic <em>Homo sapiens<\/em> have also been found, there were three crania of modern <em>Homo sapiens<\/em>. These crania, which date to between 34,000 and 10,000 years ago, were all more globular than those of archaic humans but still lower and longer than those of later modern humans (Brown 1999; Harvati 2009). When compared to one another, the crania showed significant differences from one another. Comparison of cranial measurements to other populations past and present found no connection with modern East Asians, again showing that human variation was very different from what we see today.<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 610px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image12-1.jpg\" alt=\"A cave opening amongst a dry wooded region.\" width=\"610\" height=\"458\" \/><figcaption class=\"wp-caption-text\">Figure 12.12: The entrance to the Upper Cave of the Zhoukoudian complex, where crania of three ancient modern humans were found. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Zhoukoudian_Upper_Cave.jpg\">Zhoukoudian Upper Cave<\/a> by Mutt is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><span style=\"background-color: #ff99cc\"><em>Summary of Modern <\/em>H. sapiens<em> in the Middle East and Asia<\/em><\/span><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">As in Africa, the finds of the Middle East have shown that humans were biologically diverse and had complex relationships with their environment. Work in the Levant showed an initial expansion north from the Sinai Peninsula that did not last. Away from the Levant, expansion continued. Local resources were used to make lithics and decorative items.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">The early Asian presence of modern <em>Homo sapiens<\/em> was complex and varied as befitting the massive continent. What the evidence shows is that people adapted to a wide array of environments that were far removed from Africa. From the Levant to China, humans with modern anatomy used caves that preserved signs of their presence. Faunal and floral remains found in these shelters speak to the flexibility of the human omnivorous diet as local wildlife and foliage became nourishment. Decorative items, often found as burial goods in planned graves, show a flourishing cultural life.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">Eventually, modern humans at the southeastern fringe of the geographical range of the species found their way southeast until some became the first humans in Australia.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong>Crossing to Australia<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Expansion of the first modern human Asians, still following the coast, eventually entered an area that researchers call <strong>Sunda<\/strong> before continuing on to modern Australia. Sunda was a landmass made up of the modern-day Malay Peninsula, Sumatra, Java, and Borneo. Lowered sea levels connected these places with land bridges, making them easier to traverse. Proceeding past Sunda meant navigating <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1773\">Wallacea<\/a><\/strong>, the archipelago that includes the Indonesian islands east of Borneo. In the distant past, there were many <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_864\">megafauna<\/a><\/strong>, large animals that migrating humans would have used for food and materials (such as utilizing animals\u2019 hides and bones). Further southeast was another landmass called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1774\">Sahul<\/a><\/strong>, which included New Guinea and Australia as one contiguous continent. Based on fossil evidence, this land had never seen hominins or any other primates before modern <em>Homo sapiens<\/em> arrived. Sites along this path offer clues about how our species handled the new environment to live successfully as foragers.<\/p>\n<figure style=\"width: 380px\" class=\"wp-caption alignright\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image18-1-1.jpg\" alt=\"A cranium showing a diagonal sloping forehead.\" width=\"380\" height=\"252\" \/><figcaption class=\"wp-caption-text\">Figure 12.13: Replica of the Kow Swamp 1 cranium. The shape of the braincase could be due to artificial cranial modification. A competing hypothesis is that it reflects the primitive shape of Homo erectus. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Kow_Swamp1-Homo_sapiens.jpg\">Kow Swamp1-Homo sapiens<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/14405058@N08\">Ryan Somma<\/a> from Occoquan, USA, under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/legalcode\">CC BY-SA 2.0 License<\/a> has been modified (background cleaned and color modified) and is available here under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The skeletal remains at Lake Mungo, land traditionally owned by Mutthi Mutthi, Ngiampaa, and Paakantji peoples, are the oldest known in the continent. The now-dry lake was one of a series located along the southern coast of Australia in New South Wales, far from where the first people entered from the north (Barbetti and Allen 1972; Bowler et al. 1970). Two individuals dating to around 40,000 years ago show signs of artistic and symbolic behavior, including intentional burial. The bones of Lake Mungo 1 (LM1), an adult female, were crushed repeatedly, colored with red ochre, and cremated (Bowler et al. 1970). Lake Mungo 3 (LM3), a tall, older male with a gracile cranium but robust postcranial bones, had his fingers interlocked over his pelvic region (Brown 2000).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Kow Swamp, within traditional Yorta Yorta land also in southern Australia, contained human crania that looked distinctly different from the ones at Lake Mungo (Durband 2014; Thorne and Macumber 1972). The crania, dated between 9,000 and 20,000 years ago, had extremely robust brow ridges and thick bone walls, but these were paired with globular features on the braincase (Figure 12.13).<\/p>\n<p class=\"import-Normal\">While no fossil humans have been found at the Madjedbebe rock shelter in the North Territory of Australia, more than 10,000 artifacts found there show both behavioral modernity and variability (Clarkson et al. 2017). They include a diverse array of stone tools and different shades of ochre for rock art, including mica-based reflective pigment (similar to glitter). These impressive artifacts are as far back as 56,000 years old, providing the date for the earliest-known presence of humans in Australia.<\/p>\n<h4 class=\"import-Normal\"><span style=\"background-color: #ff99cc\"><em>Summary of Modern <\/em>H. sapiens<em> in Australia<\/em><\/span><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">The overall view of the first modern humans in Australia from a biological perspective shows a high amount of skeletal diversity. This is similar to the trends seen earlier in Africa, the Middle East, and East Asia. The earliest-known arrivals brought with them a multifaceted suite of cultural practices as seen in their material culture.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong>From the Levant to Europe<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The first modern human expansion into Europe occurred after other members of our species settled East Asia and Australia. As the evidence from the Levant suggests, modern human movement to Europe may have been hampered by the presence of Neanderthals. Another obstacle was that the colder climate was incompatible with the biology of African modern <em>Homo sapiens<\/em>, <span style=\"background-color: #ffff00\">which was adapted for exposure to high temperature and ultraviolet radiation.<\/span> Still, by 40,000 years ago, modern <em>Homo sapiens<\/em> had a detectable presence. This time was also the start of the Later Stone Age or <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1775\">Upper Paleolithic<\/a><\/strong>, when there was an expansion in cultural complexity. There is a wealth of evidence from this region due to a Western bias in research, the proximity of these findings to Western scientific institutions, and the desire of Western scientists to explore their own past. <span style=\"background-color: #ff99cc\">This section will cover key evidence of early modern human life in Europe, and the typologies used to view cultural changes in this region.<\/span><\/p>\n<figure style=\"width: 323px\" class=\"wp-caption alignleft\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image11-3.jpg\" alt=\"Robust cranium with a gradually sloping forehead.\" width=\"323\" height=\"323\" \/><figcaption class=\"wp-caption-text\">Figure 12.14: This side view of the Oase 2 cranium shows the reduced brow ridges but also occipital bunning that is a sign that modern Homo sapiens interbred with Neanderthals. Credit: <a href=\"https:\/\/humanorigins.si.edu\/evidence\/human-fossils\/fossils\/oase-2\">Oase 2<\/a> by James Di Loreto &amp; Donald H. Hurlbert, <a href=\"https:\/\/www.si.edu\/\">Smithsonian<\/a> [exhibit: Human Evolution Evidence, Human Fossils] has been modified (sharpened) and <a href=\"https:\/\/www.si.edu\/termsofuse\">is used for educational and non-commercial purposes as outlined by the Smithsonian.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In Romania, the site of Pe\u0219tera cu Oase (Cave of Bones) had the oldest-known remains of modern <em>Homo sapiens<\/em> in Europe, dated to around 40,000 years ago (Trinkaus et al. 2003a). Among the bones and teeth of many animals were the fragmented cranium of one person and the mandible of another (the two bones did not fit each other). Both bones have modern human traits similar to the fossils from the Middle East, but they also had Neanderthal traits. Oase 1, the mandible, had a mental eminence but also extremely large molars (Trinkaus et al. 2003b). This mandible has yielded DNA that surprisingly is equally similar to DNA from present-day Europeans and Asians (Fu et al. 2015). This means that Oase 1 was not the direct ancestor of modern Europeans. The Oase 2 cranium has the derived traits of reduced brow ridges along with archaic wide zygomatic cheekbones and an occipital bun (Figure 12.14; Rougier et al. 2007).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Dating to around 26,000 years ago, P\u0159edmost\u00ed near P\u0159erov in the Czech Republic was a site where people buried over 30 individuals along with many artifacts. Eighteen individuals were found in one mass burial area, a few covered by the scapulae of woolly mammoths (Germonpr\u00e9, L\u00e1zni\u010dkov\u00e1-Galetov\u00e1, and Sablin 2012). The P\u0159edmost\u00ed crania were more globular than those of archaic humans but tended to be longer and lower than in later modern humans (Figure 12.15; Velem\u00ednsk\u00e1 et al. 2008). The height of the face was in line with modern residents of Central Europe. There was also skeletal evidence of dog domestication, such as the presence of dog skulls with shorter snouts than in wild wolves (Germonpr\u00e9, L\u00e1zni\u010dkov\u00e1-Galetov\u00e1, and Sablin et al. 2012). In total, P\u0159edmost\u00ed could have been a settlement dependent on mammoths for subsistence and the artificial selection of early domesticated dogs.<\/p>\n<figure style=\"width: 423px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image25-3.png\" alt=\"Black-and-white photograph of a human skull with labeled cranial landmarks.\" width=\"423\" height=\"389\" \/><figcaption class=\"wp-caption-text\">Figure 12.15: This illustration is based upon one of the surviving photographic negatives since the original fossil was lost in World War II. The modern human chin is prominent, as is an archaic occipital bun. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:P%C5%99edmost%C3%AD_9.png\">P\u0159edmost\u00ed 9<\/a> by J. Matiegka (1862\u20131941) has been modified (sharpened) and is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The sequence of modern <em>Homo sapiens<\/em> technological change in the Later Stone Age has been thoroughly dated and labeled by researchers working in Europe. Among them, the Gravettian tradition of 33,000 years to 21,000 years ago is associated with most of the known curvy female figurines, often assumed to be \u201cVenus\u201d figures. Hunting technology also advanced in this time with the first known boomerang, <strong>atlatl<\/strong> (spear thrower), and archery. The Magdalenian tradition spread from 17,000 to 12,000 years ago. This culture further expanded on fine bone tool work, including barbed spearheads and fishhooks (Figure 12.16).<\/p>\n<figure style=\"width: 511px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image2-1-1.jpg\" alt=\"Long, thin spear tips. Many have barbs, others are smooth.\" width=\"511\" height=\"494\" \/><figcaption class=\"wp-caption-text\">Figure 12.16: This drawing from 1891 shows an array of Magdalenian-style barbed points found in the burial of a reindeer hunter. They were carved from antler. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:La_station_quaternaire_de_Raymonden_(...)Hardy_Michel_bpt6k5567846s_(2).jpg\">La station quaternaire de Raymonden (&#8230;)Hardy Michel bpt6k5567846s (2)<\/a> by M. F\u00e9auxis, original by Michel Hardy (1891), is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Among the many European sites dating to the Later Stone Age, the famous cave art sites deserve mention. Chauvet-Pont-d&#8217;Arc Cave in southern France dates to separate Aurignacian occupations 31,000 years ago and 26,000 years ago. Over a hundred art pieces representing 13 animal species are preserved, from commonly depicted deer and horses to rarer rhinos and owls. Another French cave with art is Lascaux, which is several thousand years younger at 17,000 years ago in the Magdalenian period. At this site, there are over 6,000 painted figures on the walls and ceiling (Figure 12.17). Scaffolding and lighting must have been used to make the paintings on the walls and ceiling deep in the cave. Overall, visiting Lascaux as a contemporary must have been an awesome experience: trekking deeper in the cave lit only by torches giving glimpses of animals all around as mysterious sounds echoed through the galleries.<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 605px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image24-2-1.jpg\" alt=\"Charcoal painting of a bull seen from the side.\" width=\"605\" height=\"454\" \/><figcaption class=\"wp-caption-text\">Figure 12.17: Photograph of just one surface with cave art at Lascaux Cave. The most prominent piece here is the Second Bull, found in a chamber called the Hall of Bulls. Smaller cattle and horses are also visible. Credit: <a href=\"https:\/\/whc.unesco.org\/en\/documents\/108435\">Lascaux cave (document 108435) Prehitoric Sites and Decorated Caves of the V\u00e9z\u00e8re Valley (France)<\/a> by Francesco Bandarin, <a href=\"https:\/\/whc.unesco.org\/\">\u00a9 UNESCO<\/a>, has been modified (color modified) and is under a <a href=\"https:\/\/whc.unesco.org\/en\/licenses\/6\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><span style=\"background-color: #ff99cc\"><em>Summary of Modern <\/em>H. sapiens<em> in Europe<\/em><\/span><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">Study of Europe in the Upper Paleolithic gives a more detailed view of the general pattern of biological and cultural change linked with the arrival of modern <em>Homo sapiens<\/em>. The modern humans experienced a rapidly changing culture that went through waves of complexity and refinement. Skeletally, the increasing globularity of the cranium and the gracility of the rest of the skeleton continued, though with unique regional traits, too. The cave art sites showed a deeper exploration of creativity though the exact meaning is unclear. With survival dependent on the surrounding ecology, painting the figures may have connected people to important and impressive wildlife at both a physical and spiritual level. Both reverence for animals and the use of caves for an enhanced sensory experience are common to cultures past and present.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong>Peopling of the Americas<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">By 25,000 years ago, our species was the only member of <em>Homo<\/em> left on Earth. Gone were the Neanderthals, Denisovans, <em>Homo naledi,<\/em> and <em>Homo floresiensis<\/em>. The range of modern <em>Homo sapiens<\/em> kept expanding eastward into\u2014using the name given to this area by Europeans much later\u2014the Western Hemisphere. This section will address what we know about the peopling of the Americas, from the first entry to these continents to the rapid spread of Indigenous Americans across its varied environments.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">While evidence points to an ancient land bridge called <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1778\"><strong>Beringia<\/strong><\/a> that allowed people to cross from what is now northeastern Siberia into modern-day Alaska, what people did to cross this land bridge is still being investigated. For most of the 20th century, the accepted theory was the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1779\"><strong>Ice-Free Corridor model<\/strong><\/a>. It stated that northeast Asians (East Asians and Siberians) first expanded across Beringia inland through a passage between glaciers that opened into the western Great Plains of the United States, just east of the Rocky Mountains, around 13,000 years ago (Swisher et al. 2013). While life up north in the cold environment would have been harsh, migrating birds and an emerging forest might have provided sustenance as generations expanded through this land (Potter et al. 2018).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">However, in recent decades, researchers have accumulated evidence against the Ice-Free Corridor model. Archaeologist K. R. Fladmark (1979) brought the alternate <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1780\"><strong>Coastal Route model<\/strong><\/a> into the archaeological spotlight; researcher Jon M. Erlandson has been at the forefront of compiling support for this theory (Erlandson et al. 2015). The new focus is the southern edge of the land bridge instead of its center: About 16,000 years ago, members of our species expanded along the coastline from northeast Asia, east through Beringia, and south down the Pacific Coast of North America while the inland was still sealed off by ice. The coast would have been free of ice at least part of the year, and many resources would have been found there, such as fish (e.g., salmon), mammals (e.g., whales, seals, and otters), and plants (e.g., seaweed).<\/p>\n<h4 class=\"import-Normal\"><em>South through the Americas<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">When the first modern <em>Homo sapiens<\/em> reached the Western Hemisphere, the spread through the Americas was rapid. Multiple migration waves crossed from North to South America (Posth et al. 2018). Our species took advantage of the lack of hominin competition and the bountiful resources both along the coasts and inland. The Americas had their own wide array of megafauna, which included woolly mammoths (Figure 12.18), mastodons, camels, horses, ground sloths, giant tortoises, and\u2014a favorite of researchers\u2014a two-meter-tall beaver. The reason we cannot see these amazing animals today may be that resources gained from these fauna were crucial to the survival for people over 12,000 years ago (Araujo et al. 2017). Several sites are notable for what they add to our understanding of the distant past in the Americas, including interactions with megafauna and other elements of the environment.<\/p>\n<figure style=\"width: 242px\" class=\"wp-caption alignleft\"><img loading=\"lazy\" decoding=\"async\" class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image3-2-1.jpg\" alt=\"A mammoth model with long curving tusks.\" width=\"242\" height=\"323\" \/><figcaption class=\"wp-caption-text\">Figure 12.18: Life-size reconstruction of a woolly mammoth at the Page Museum, part of the La Brea Tar Pits complex in Los Angeles, California. Outside of Africa, megafauna such as this went extinct around the time that humans entered their range. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-14\/\">Woolly Mammoth<\/a> (at <a href=\"https:\/\/tarpits.org\/\">La Brea Tar Pits &amp; Museum<\/a>) by Keith Chan is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\"><span style=\"background-color: #ff9900\">A 2019 discovery may allow researchers to improve theories about the peopling of the Americas. In White Sands National Park, New Mexico, 60 human footprints have been astonishingly dated to around 22,000 years ago (Bennett et al. 2021). This date and location do not match either the Ice-Free Corridor or Coastal Route models. Researchers are now working to verify the find and adjust previous models to account for the new evidence. This groundbreaking find is sparking new theories; it is another example of the fast pace of research performed on our past.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Monte Verde is a landmark site that shows that the human population had expanded down the whole vertical stretch of the Americas to Chile by 14,600 years ago, <span style=\"background-color: #ffff00\">only a few thousand years after humans first entered the Western Hemisphere from Alaska.<\/span> The site has been excavated by archaeologist Tom D. Dillehay and his team (2015). The remains of nine distinct edible species of seaweed at the site shows familiarity with coastal resources and relates to the Coastal Route model by showing a connection between the inland people and the sea.<\/p>\n<figure style=\"width: 254px\" class=\"wp-caption alignright\"><img loading=\"lazy\" decoding=\"async\" class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image21-4.png\" alt=\"A long stone point with small chips around the edge.\" width=\"254\" height=\"362\" \/><figcaption class=\"wp-caption-text\">Figure 12.19: The Clovis point has a distinctive structure. It has a wide tip, and its base has two small projections. This example was carved from chert and found in north-central Ohio, dated to around 11,000 years ago. Credit: <a href=\"https:\/\/www.si.edu\/object\/chndm_15.2012.25\">Clovis Point<\/a> (15.2012.25) by <a href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [Department of Anthropology; Cooper Hewitt, Smithsonian Design Museum] <a href=\"https:\/\/www.si.edu\/termsofuse\">is used for educational and non-commercial purposes as outlined by the Smithsonian.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Named after the town in New Mexico, the Clovis stone-tool style is the first example of a widespread culture across much of North America, between 13,400 and 12,700 years ago (Miller, Holliday, and Bright 2013). Clovis points were fluted with two small projections, one on each end of the base, facing away from the head (Figure 12.19). The stone points found at this site match those found as far as the Canadian border and northern Mexico, and from the west coast to the east coast of the United States. Fourteen Clovis sites also contained the remains of mammoths or mastodons, suggesting that hunting megafauna with these points was an important part of life for the Clovis people. After the spread of the Clovis style, it diversified into several regional styles, keeping some of the Clovis form but also developing their own unique touches.<\/p>\n<p><span style=\"text-decoration: underline;background-color: #ff99cc\">(maybe inlcude a special topic\/dig deeper from Dr.Steeves talking about Clovis culture and effects on Indigenous histories)<\/span><\/p>\n<h4 class=\"import-Normal\"><span style=\"background-color: #ff99cc\"><em>Summary of Modern <\/em>H. sapiens<em> in the Americas<\/em><\/span><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">Research in Native American origins found some surprising details, refining older models. Genetically, the migration can be considered one long period of movement with splits into regional populations. This finding matches the sudden appearance and diversification of the homegrown Clovis culture. A few thousand years after arrival into the hemisphere, people had already covered the Americas from north to south.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">The peopling of the Americas also had a lot of elements in common with the prior spread of humans across Africa, Europe, Asia, and Australia. In all of these expansions, these pioneers explored new lands that tested their ability to adapt, both culturally and biologically. Besides stone-tool technology, the use of ochre as decoration was seen from South Africa to South America. The coasts and rivers were likely avenues in the movement of people, artifacts, and ideas, outlining the land masses while providing access to varied environments. The presence of megafauna aided human success, but this resource was eventually depleted in many parts of the world.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong>The Big Picture: The Assimilation Hypothesis<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">How do researchers make sense of all of these modern <em>Homo sapiens<\/em> discoveries that cover over 300,000 years of time and stretch across every continent except Antarctica? How was modern <em>Homo sapiens<\/em> related to archaic <em>Homo sapiens<\/em>?<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1782\">Assimilation hypothesis<\/a><\/strong> proposes that modern <em>Homo sapiens<\/em> evolved in Africa first and expanded out but also interbred with the archaic <em>Homo sapiens<\/em> they encountered outside Africa (Figure 12.20). This hypothesis is powerful since it explains why Africa has the oldest modern human fossils, why early modern humans found in Europe and Asia bear a resemblance to the regional archaics, and why traces of archaic DNA can be found in our genomes today (Dannemann and Racimo 2018; Reich et al. 2010; Reich et al. 2011; Slatkin and Racimo 2016; Smith et al. 2017; Wall and Yoshihara Caldeira Brandt 2016).<\/p>\n<figure style=\"width: 443px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image28-2.png\" alt=\"African Homo erectus expands and gives rise to archaics and modern Homo sapiens groups.\" width=\"443\" height=\"471\" \/><figcaption class=\"wp-caption-text\">Figure 12.20: This diagram shows archaic humans, having evolved from Homo erectus, expanded from Africa and established the Neanderthal and Denisovan groups. In Africa, archaic humans evolved modern traits and expanded from the continent as well, interbreeding with two archaic groups across Europe and Asia. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-14\/\">Assimilation Model (Figure 12.23)l<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Keith Chan and Katie Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">While researchers have produced a model that satisfies the data, there are still a lot of questions for paleoanthropologists to answer regarding our origins. What were the patterns of migration in each part of the world? Why did the archaic humans go extinct? In what ways did archaic and modern humans interact? The definitive explanation of how our species started and what our ancestors did is still out there to be found. You are now in a great place to welcome the next discovery about our distant past\u2014maybe you\u2019ll even contribute to our understanding as well.<\/p>\n<h2 class=\"import-Normal\">The Chain Reaction of Agriculture<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ccffcc\">While it may be hard to imagine today, for most of our species\u2019 existence we were nomadic: moving through the landscape without a singular home. Instead of a refrigerator or pantry stocked with food, we procured nutrition and other resources as needed based on what was available in the environment.<\/span> <span style=\"background-color: #ffff00\">Instead of collecting and displaying shelves of stuff, we kept our possessions small for mobility.<\/span> <span style=\"background-color: #ff99cc\">This section gives an overview of how the foraging lifestyle enabled the expansion of our species and how the invention of a new way of life caused a chain reaction of cultural change.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong>The Foraging Tradition<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">There are a variety of possible <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1783\">subsistence strategies<\/a><\/strong>, or methods of finding sustenance and resources. To understand our species is to understand the subsistence strategy of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1114\">foraging<\/a><\/strong>, or the search for resources in the environment. While most (but not all) humans today live in cultures that practice <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1784\">agriculture<\/a> <\/strong>(whereby we greatly shape the environment to mass produce what we need), we have spent far more time as nomadic foragers than as settled agriculturalists. As such, <span style=\"background-color: #ffff00\">our traits have evolved to be primarily geared toward foraging. For instance, our efficient bipedalism allows persistence-hunting across long distances as well as movement from resource to resource.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">How does human foraging, also known as hunting and gathering, work? Anthropologists have used all four fields to answer this question (see Ember n.d.). Typically, people formed <strong>bands<\/strong>, or kin-based groups of around 50 people or less (rarely over 100). A band\u2019s organization would be <strong>e<\/strong><strong>galitarian<\/strong>, with a flexible hierarchy based on an individual\u2019s age, level of experience, and relationship with others. Everyone would have a general knowledge of the skills assigned to their gender roles, rather than specializing in different occupations. A band would be able to move from place to place in the environment, using knowledge of the area to forage (Figure 12.21). In varied environments\u2014from savannas to tropical forests, deserts, coasts, and the Arctic circle\u2014people found sustenance needed for survival. <span style=\"background-color: #ffff00\">Our species\u2019s omnivorousness and cultural abilities led us to excel in the generalist-specialist niche.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 565px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image22.jpg\" alt=\"A hunter holding a bow is crouched among dry grass.\" width=\"565\" height=\"377\" \/><figcaption class=\"wp-caption-text\">Figure 12.21: A present-day San man in Namibia demonstrates hunting using archery. Anthropologists study the San today to learn about the persistence of foraging as a viable lifestyle, while noting how these cultures have changed over time and how they interact with other groups. Credit: <a href=\"https:\/\/www.flickr.com\/photos\/charlesfred\/2129551464\">San hunter w\u0131th bow and arrow<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/charlesfred\/\">CharlesFred<\/a> has been modified (color modified) and is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ffff00\">Humans made extensive use of the foraging subsistence strategy, but this lifestyle did have limitations. The ease of foraging depended on the richness of the environment. Due to the lack of storage, resources had to be dependably found when needed. While a bountiful environment would require just a few hours of foraging a day and could lead to a focus on one location, the level and duration of labor increased greatly in poor or unreliable environments.<\/span> Labor was also needed to process the acquired resources, which contributed to the foragers\u2019 daily schedule (Crittenden and Schnorr 2017).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ffff00\">The adaptations to foraging found in modern <em>Homo sapiens<\/em> may explain why our species became so successful both within Africa and in the rapid expansion around the world. Overcoming the limitations, each generation at the edge of our species\u2019s range would have found it beneficial to expand a little further, keeping contact with other bands but moving into unexplored territory where resources were more plentiful. The cumulative effect would have been the spread of modern <em>Homo sapiens<\/em> across continents and hemispheres.<\/span><\/p>\n<h2 class=\"import-Normal\"><strong>Why Agriculture?<\/strong><\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">After hundreds of thousands of years of foraging, some groups of people around 12,000 years ago started to practice agriculture. This transition, called the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1787\"><strong>Neolithic Revolution<\/strong>,<\/a> occurred at the start of the <strong>Holocene<\/strong> epoch. While the reasons for this global change are still being investigated, two likely co-occurring causes are a growing human population and natural global climate change.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Overcrowding could have affected the success of foraging in the environment, leading to the development of a more productive subsistence strategy (Cohen 1977). Foraging works best with low population densities since each band needs a lot of space to support itself. If too many people occupy the same environment, they deplete the area faster. The high population could exceed the <strong>carrying capacity<\/strong>, or number of people a location can reliably support. Reaching carrying capacity on a global level due to growing population and limited areas of expansion would have been an increasingly pressing issue after the expansion through the major continents by 14,600 years ago.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">A changing global climate immediately preceded the transition to agriculture, so researchers have also explored a connection between the two events. Since the <strong>Last Glacial Maximum<\/strong> of 23,000 years ago, the Earth slowly warmed. Then, from 13,000 to 11,700 years ago, the temperature in most of the Northern Hemisphere dropped suddenly in a phenomenon called the <strong>Younger Dryas<\/strong>. Glaciers returned in Europe, Asia, and North America. In Mesopotamia, which includes the Levant, the climate changed from warm and humid to cool and dry. The change would have occurred over decades, disrupting the usual nomadic patterns and subsistence of foragers around the world. The disruption to foragers due to the temperature shift could have been a factor in spurring a transition to agriculture. Researchers Gregory K. Dow and colleagues (2009) believe that foraging bands would have clustered in the new resource-rich places where people started to direct their labor to farming the limited area. After the Younger Dryas ended, people expanded out of the clusters with their agricultural knowledge (Figure 12.22).<\/p>\n<figure style=\"width: 570px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image7-6.png\" alt=\"Map shows that agriculture was invented in at least six parts of the world.\" width=\"570\" height=\"267\" \/><figcaption class=\"wp-caption-text\">Figure 12.22: The map shows the areas where agriculture was independently invented around the world and where they spread. Blue arrows show the spread of agriculture from these zones to other regions. <a href=\"https:\/\/docs.google.com\/document\/d\/1VUDKMBJYS_jNONjLxT04jQN0_z9Ua50BRN6auGSHUuU\/edit\">A full text description of this image is available<\/a>. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Centres_of_origin_and_spread_of_agriculture.svg\">Centres of origin and spread of agriculture<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Joe_Roe\">Joe Roe<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The double threat of the limitation of human continental expansion and the sudden global climate change may have placed bands in peril as more populations outpaced their environment\u2019s carrying capacity. <span style=\"background-color: #ffff00\">Not only had a growing population led to increased competition with other bands, but environments worldwide shifted to create more uncertainty. As people in different areas around the world faced this unpredictable situation, they became the independent inventors of agriculture.<\/span><\/p>\n<h2 class=\"import-Normal\"><strong>Agriculture around the World<\/strong><\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Due to global changes to the human experience starting from 12,000 years ago, <span style=\"background-color: #ffff00\">cultures with no knowledge of each other turned toward intensely farming their local resources<\/span> (see Figure 12.22). <span style=\"background-color: #ffff00\">The first farmers engaged in artificial selection of their domesticates to enhance useful traits over generations. The switch to agriculture took time and effort with no guarantee of success and constant challenges (e.g. fires, droughts, diseases, and pests).<\/span> The regions with the most widespread impact in the face of these obstacles became the primary centers of agriculture (Figure 12.23; Fuller 2010):<\/p>\n<ul>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Mesopotamia: The Fertile Crescent from the Tigris and Euphrates rivers through the Levant was where bands started to domesticate plants and animals around 12,000 years ago. The connection between the development of agriculture and the Younger Dryas was especially strong here. Farmed crops included wheat, barley, peas, and lentils. This was also where cattle, pigs, sheep, and goats were domesticated.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">South and East Asia: Multiple regions across this land had varieties of rice, millet, and soybeans by 10,000 years ago. Pigs were farmed with no connection to Mesopotamia. Chickens were also originally from this region, bred for fighting first and food second.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">New Guinea: Agriculture started here 10,000 years ago. Bananas, sugarcane, and taro were native to this island. Sweet potatoes were brought back from voyages to South America around the year C.E. 1000. No known animal farming occurred here.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Mesoamerica: Agriculture from Central Mexico to northern South America also occurred from 10,000 years ago; it was also only plant based. Maize was a crop bred from teosinte grass, which has become one of the global staples. Beans, squash, and avocados were also grown in this region.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">The Andes: Starting around 8,000 years ago, local domesticated plants started with squash but later included potatoes, tomatoes, beans, and quinoa. Maize was brought down from Mesoamerica. The main farm animals were llamas, alpacas, and guinea pigs.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Sub-Saharan Africa: This region went through a change 5,000 years ago called the Bantu expansion. The Bantu agriculturalists were established in West Central Africa and then expanded south and east. Native varieties of rice, yams, millet, and sorghum were grown across this area. Cattle were also domesticated here.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Eastern North America: This region was the last major independent agriculture center, from 4,000 years ago. Squash and sunflower are the produce from this region that are most known today, though sumpweed and pitseed goosefoot were also farmed. Hunting was still the main source of animal products.<\/li>\n<\/ul>\n<figure style=\"width: 482px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image23-1-1.jpg\" alt=\"Farmers plow a flooded field. Each plow is pulled by two oxen.\" width=\"482\" height=\"320\" \/><figcaption class=\"wp-caption-text\">Figure 12.23: Rice farmers in the present day using draft cattle to prepare their field. Credit: <a href=\"https:\/\/www.flickr.com\/photos\/ricephotos\/7554483250\">Plowing muddy field using cattle<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/ricephotos\/\">IRRI Photos<\/a> (International Rice Research Institute) has been modified (color modified) and is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">By 5,000 years ago, our species was well within the Neolithic Revolution. Agriculturalists spread to neighboring parts of the world with their domesticates, further expanding the use of this subsistence strategy. <span style=\"background-color: #ffff00\">From this point, the human species changed from being primarily foragers to primarily agriculturalists with skilled control of their environments.<\/span> The planet changed from mostly unaffected by human presence to being greatly transformed by humans. The revolution took millennia, but it was a true revolution as our species\u2019 lifestyle was dramatically reshaped.<\/p>\n<h3 class=\"import-Normal\"><strong>Cultural Effects of Agriculture<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The worldwide adoption of agriculture altered the course of human culture and history forever. The core change in human culture due to agriculture is the move toward not moving: rather than live a nomadic lifestyle, farmers had to remain in one area to tend to their crops and livestock. The term for living bound to a certain location is <strong>sedentarism<\/strong>. <span style=\"background-color: #ffff00\">This led to new aspects of life that were uncommon among foragers: the construction of permanent shelters and agricultural infrastructure, such as fields and irrigation, plus the development of storage technology, such as pottery, to preserve extra resources in case of future instability.<\/span><\/p>\n<figure style=\"width: 359px\" class=\"wp-caption alignright\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image20-1-1.jpg\" alt=\"Multistory buildings surrounding a greek-style plaza.\" width=\"359\" height=\"270\" \/><figcaption class=\"wp-caption-text\">Figure 12.24: View of downtown San Diego taken by the author at a shopping complex during a break from jury duty. Here, people live amongst structures that facilitate commerce, government, tourism, and art. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-14\/\">Downtown San Diego (October 13, 2016; Figure 12.28)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Keith Chan is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The high productivity of successful agriculture sparked further changes (Smith 2009). <span style=\"background-color: #ffff00\">Since successful agriculture produced a much-greater amount of food and other resources per unit of land compared to foraging, the population growth rate skyrocketed.<\/span> <span style=\"background-color: #ccffcc\">The surplus of a bountiful harvest also provided insurance for harder times, reducing the risk of famine. Changes happened to society as well. With a few farming households producing enough food to feed many others, other people could focus on other tasks. So began specialization into different occupations such as craftspeople, traders, religious figures, and artists, spurring innovation in these areas as people could now devote time and effort toward specific skills. These interdependent people would settle an area together for convenience. The growth of these settlements led to <strong>urbanization<\/strong>, the founding of cities that became the foci of human interaction (Figure 12.24).<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The formation of cities led to new issues that sparked the growth of further specializations, called <strong>institutions<\/strong>. These are cultural constructs that exist beyond the individual and have wide control over a population. Leadership of these cities became hierarchical with different levels of rank and control. The stratification of society increased social inequality between those with more or less power over others. Under leadership, people built impressive <strong>monumental architecture<\/strong>, such as pyramids and palaces, that embodied the wealth and power of these early cities. Alliances could unite cities, forming the earliest states. In several regions of the world, state organization expanded into empires, wide-ranging political entities that covered a variety of cultures.<\/p>\n<p><span style=\"text-decoration: underline;background-color: #00ffff\">(Inlcude Special Topic about the Haudesaunee\/Iroquois confederacy)<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Urbanization brought new challenges as well. The concentration of sedentary peoples was ideal for infectious diseases to thrive since they could jump from person to person and even from livestock to person (Armelagos, Brown, and Turner 2005). While successful agriculture provided a large surplus of food to thwart famine, the food produced offered less diverse food sources than foragers\u2019 diets (Cohen and Armelagos 1984; Cohen and Crane-Kramer 2007). This shift in nutrition caused other diseases to flourish among those who adopted farming, such as dental cavities and malocclusion (the misalignment of teeth caused by soft, agricultural diets). The need to extract \u201cwisdom teeth\u201d or third molars seen in agricultural cultures today stems from this misalignment between the environment our ancestors adapted to and our lifestyles today.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">As the new disease trends show, the adoption of agriculture and the ensuing cultural changes were not entirely positive. It is also important to note that this is not an absolutely linear progression of human culture from simple to complex. In many cases, empires have collapsed and, in some cases, cities dispersed to low-density bands that rejected institutions. However, a global trend has emerged since the adoption of agriculture, wherein population and social inequality have increased, leading to the massive and influential nation-states of today.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The rise of states in Europe has a direct impact on many of this book\u2019s topics. Science started as a European cultural practice by the upper class that became a standardized way to study the world. Education became an institution to provide a standardized path toward producing and gaining knowledge. The scientific study of human diversity, embroiled in the race concept that still haunts us today, was connected to the European slave trade and colonialism.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Also starting in Europe, the Industrial Revolution of the 19th century turned cities into centers of mass manufacturing and spurred the rapid development of inventions (Figure 12.25). In the technologically interconnected world of today, human society has reached a new level of complexity with <strong>globalization<\/strong>. In this system, goods are mass-produced and consumed in different parts of the world, weakening the reliance on local farms and factories. The imbalanced relationship between consumers and producers of goods further increases economic inequality.<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 465px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image1-3.jpg\" alt=\"A yellow farm vehicle driving into crops in a field.\" width=\"465\" height=\"310\" \/><figcaption class=\"wp-caption-text\">Figure 12.25: This combine harvester can collect and process grain at a massive scale. Our food now commonly comes from enormous farms located around the world. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Combine_CR9060.jpeg\">Combine CR9060<\/a> by Hertzsprung is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">As states based on agriculture and industry keep exerting influence on humanity today, there are people, like the Hadzabe of Tanzania, who continue to live a lifestyle centered on foraging. Due to the overwhelming force that agricultural societies exert, foragers today have been marginalized to live in the least habitable parts of the world\u2014the areas that are not conducive to farming, such as tropical rainforests, deserts, and the Arctic (Headland et al. 1989). Foragers can no longer live in the abundant environments that humans would have enjoyed before the Neolithic Revolution. Interactions with agriculturalists are typically imbalanced, with trade and other exchanges heavily favoring the larger group. One of anthropology\u2019s important roles today is to intelligently and humanely manage equitable interactions between people of different backgrounds and levels of influence.<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Indigenous Land Management<\/h2>\n<p class=\"import-Normal\">Insight into the lives of past modern humans has evolved as researchers revise previous theories and establish new connections with Indigenous knowledge holders.<\/p>\n<p class=\"import-Normal\">The outdated view of foraging held that people lived off of the land without leaving an impact on the environment. Accompanying this idea was anthropologist Marshall Sahlins\u2019s (1968) proposal that foragers were the \u201coriginal affluent society\u201d since they were meeting basic needs and achieving satisfaction with less work hours than agriculturalists and city-dwellers. This view countered an earlier idea that foragers were always on the brink of starvation. Sahlins\u2019s theory took hold in the public eye as an attractive counterpoint to our busy contemporary lives in which we strive to meet our endless wants.<\/p>\n<p class=\"import-Normal\">A fruitful type of study involving researchers collaborating with Indigenous experts has found that foragers did not just live off the land with minimal effort nor were they barely surviving in unchanging environments. Instead, they shaped the landscape to their needs using labor and strategies that were more subtle than what European colonizers and subsequent researchers were used to seeing. Research from two regions shows the latest developments in understanding Indigenous land management.<\/p>\n<p class=\"import-Normal\">In British Columbia, Canada, the bridging of scientific and Indigenous perspectives has shown that the forests of the region are not untouched wilderness but, rather, have been crafted by Indigenous peoples thousands of years ago. Forest gardens adjacent to archaeological sites show higher plant diversity than unmanaged places even after 150 years (Armstrong et al. 2021). On the coast, 3,500-year-old archaeological sites are evidence of constructed clam gardens, according to Indigenous experts (Lepofsky et al. 2015). Another project, in consultation with Elders of the T\u2019exelc (William Lakes First Nation) in British Columbia, introduced researchers to explanations of how forests were managed before the practice was disrupted by European colonialism (Copes-Gerbitz et al. 2021). Careful management of controlled fires reduced the density of the forest to favor plants such as raspberries and allow easier movement through the landscape.<\/p>\n<p class=\"import-Normal\">Similarly, the study of landscapes in Australia, in consultation with Aboriginal Australians today, shows that areas previously considered wilderness by scientists were actually the result of controlling fauna and fires. The presence of grasslands with adjacent forests were purposely constructed to attract kangaroos for hunting (Gammage 2008). People also managed other animal and insect life, from emus to caterpillars. In Tasmania, a shift from productive grassland to wildfire-prone rainforest occurred after Aboriginal Australian land management was replaced by British colonial rule (Fletcher, Hall, and Alexander 2021). The site of Budj Bim of the Gunditjmara people has archaeological features of <strong>aquaculture<\/strong>, or the farming of fish, that date back 6,600 years (McNiven et al. 2012; McNiven et al. 2015). These examples show that Indigenous knowledge of how to manipulate the environment may be invaluable at the state level, such as by creating an Aboriginal ranger program to guide modern land management.<\/p>\n<\/div>\n<h2 class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Conclusion<\/span><\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">Modern <em>Homo sapiens<\/em> is the species that took the hominin lifestyle the furthest to become the only living member of that lineage. The largest factor that allowed us to persist while other hominins went extinct was likely our advanced ability to culturally adapt to a wide variety of environments. Our species, with its skeletal and behavioral traits, was well-suited to be generalist-specialists who successfully foraged across most of the world\u2019s environments. The biological basis of this adaptation was our reorganized brain that facilitated innovation in cultural adaptations and intelligence for leveraging our social ties and finding ways to acquire resources from the environment. As the brain\u2019s ability increased, it shaped the skull by reducing the evolutionary pressure to have large teeth and robust cranial bones to produce the modern <em>Homo sapiens<\/em> face.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">Our ability to be generalist-specialists is seen in the geographical range that modern <em>Homo sapiens<\/em> covered in 300,000 years. In Africa, our species formed from multiregional gene flow that loosely connected archaic humans across the continent. People then expanded out to the rest of the continental Eurasia and even further to the Americas.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">For most of our species\u2019s existence, foraging was the general subsistence strategy within which people specialized to culturally adapt to their local environment. With omnivorousness and mobility, people found ways to extract and process resources, shaping the environment in return. When resource uncertainty hit the species, people around the world focused on agriculture to have a firmer control of sustenance. The new strategy shifted human history toward exponential growth and innovation, leading to our high dependence on cultural adaptations today.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"background-color: #ff99cc\">While a cohesive image of our species has formed in recent years, there is still much to learn about our past. The work of many driven researchers shows that there are amazing new discoveries made all the time that refine our knowledge of human evolution. Technological innovations such as DNA analysis enable scientists to approach lingering questions from new angles. The answers we get allow us to ask even more insightful questions that will lead us to the next revelation. Like the pink limestone strata at Jebel Irhoud, previous effort has taken us so far and you are now ready to see what the next layer of discovery holds.<\/span><\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\"><del>Special Topic:<\/del> The Future of Humanity<\/h2>\n<p class=\"import-Normal\">A common question stemming from understanding human evolution is: What will the genetic and biological traits of our species be hundreds of thousands of years in the future? When faced with this question, people tend to think of directional selection. Maybe our braincases will be even larger, resembling the large-headed and small-bodied aliens of science fiction (Figure 12.26). Or, our hands could be specialized for interacting with our touch-based technology with less risk of repetitive injury. These ideas do not stand up to scrutiny. Since natural selection is based on adaptations that increase reproductive success, any directional change must be due to a higher rate of producing successful offspring compared to other alleles. Larger brains and more agile fingers would be convenient to possess, but they do not translate into an increase in the underlying allele frequencies.<\/p>\n<figure style=\"width: 571px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image8-4.png\" alt=\"One human has typical features; the other has a tall braincase.\" width=\"571\" height=\"279\" \/><figcaption class=\"wp-caption-text\">Figure 12.26: Will we evolve toward even more globular brains? Actually, this trend is not likely to continue for our species. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-14\/\">Hypothetical image of future human evolution (Figure 12.30)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Scientists are hesitant to professionally speculate on the unknowable, and we will never know what is in store for our species one thousand or one million years from now, but there are two trends in human evolution that may carry on into the future: increased genetic variation and a reduction in regional differences.<\/p>\n<p class=\"import-Normal\">Rather than a directional change, genetic variation in our species could expand. Our technology can protect us from extreme environments and pathogens, even if our biological traits are not tuned to handle these stressors. The rapid pace of technological advancement means that biological adaptations will become less and less relevant to reproductive success, so nonbeneficial genetic traits will be more likely to remain in the gene pool. Biological anthropologist Jay T. Stock (2008) views environmental stress as needing to defeat two layers of protection before affecting our genetics. The first layer is our cultural adaptations. Our technology and knowledge can reduce pressure on one\u2019s genotype to be \u201cjust right\u201d to pass to the next generation. The second defense is our flexible physiology, such as our acclimatory responses. Only stressors not handled by these powerful responses would then cause natural selection on our alleles. These shields are already substantial, and cultural adaptations will only keep increasing in strength.<\/p>\n<p class=\"import-Normal\">The increasing ability to travel far from one\u2019s home region means that there will be a mixing of genetic variation on a global level in the future of our species. In recent centuries, gene flow of people around the world has increased, creating admixture in populations that had been separated for tens of thousands of years. For skin color, this means that populations all around the world could exhibit the whole range of skin colors, rather than the current pattern of decreasing melanin pigment farther from the equator. The same trend of intermixing would apply to all other traits, such as blood types. While our genetics will become more varied, the variation will be more intermixed instead of regionally isolated.<\/p>\n<p class=\"import-Normal\">Our distant descendants will not likely be dextrous ultraintellectuals; more likely, they will be a highly variable and mobile species supported by novel cultural adaptations that make up for any inherited biological limitations. Technology may even enable the editing of DNA directly, changing these trends. With the uncertainty of our future, these are just the best-educated guesses for now. Our future is open and will be shaped little by little by the environment, our actions, and the actions of our descendants.<\/p>\n<\/div>\n<h2 class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Hominin Species Summary<\/span><\/h2>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Modern<em> Homo sapiens<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">315,000 years ago to present<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">Starting in Africa, then expanding around the world<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">Cro-Magnon individuals, discovered 1868 in Dordogne, France. Otzi the Ice Man, discovered 1991 in the Alps between Austria and Italy. Kennewick man, discovered 1996 in Washington state.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">1400 cc average<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Extremely small with short cusps.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">An extremely globular brain case and gracile features throughout the cranium. The mandibular symphysis forms a chin at the anterior-most point.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">Gracile skeleton adapted for efficient bipedal locomotion at the expense of the muscular strength of most other large primates.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">Extremely extensive and varied culture with many spoken and written languages. Art is ubiquitous. Technology is broad in complexity and impact on the environment.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">The only living hominin. Chimpanzees and bonobos are the closest living relatives.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Review Questions<strong><br \/>\n<\/strong><\/h2>\n<ul>\n<li>What are the skeletal and behavioral traits that define modern <em>Homo sapiens<\/em>? What are the evolutionary explanations for its presence?<\/li>\n<li>What are some creative ways that researchers have learned about the past by studying fossils and artifacts?<\/li>\n<li>How do the discoveries mentioned in \u201cFirst Africa, Then the World\u201d fit the Assimilation model?<\/li>\n<li>What is foraging? What adaptations do we have for this subsistence strategy? Could you train to be a skilled forager?<\/li>\n<li>What are aspects of your life that come from dependence on agriculture and its cultural effects? Where did the ingredients of your favorite foods originate from?<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<h2 class=\"__UNKNOWN__\">Key Terms<\/h2>\n<div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\"><strong>African multiregionalism<\/strong>: The idea that modern <em>Homo sapiens<\/em> evolved as a complex web of small regional populations with sporadic gene flow among them.<\/p>\n<p class=\"import-Normal\"><strong>Agriculture<\/strong>: The mass production of resources through farming and domestication.<\/p>\n<p class=\"import-Normal\"><strong>Aquaculture<\/strong>: The farming of fish using techniques such as trapping, channels, and artificial ponds.<\/p>\n<p class=\"import-Normal\"><strong>Assimilation <\/strong><strong>hypothesis<\/strong>: Current theory of modern human origins stating that the species evolved first in Africa and interbred with archaic humans of Europe and Asia.<\/p>\n<p class=\"import-Normal\"><strong>Atlatl<\/strong>: A handheld spear thrower that increased the force of thrown projectiles.<\/p>\n<p class=\"import-Normal\"><strong>Band<\/strong>: A small group of people living together as foragers.<\/p>\n<p class=\"import-Normal\"><strong>Beringia<\/strong>: Ancient landmass that connected Siberia and Alaska. The ancestors of Indigenous Americans would have crossed this area to reach the Americas.<\/p>\n<p class=\"import-Normal\"><strong>Carrying capacity<\/strong>: The amount of organisms that an environment can reliably support.<\/p>\n<p class=\"import-Normal\"><strong>Coastal Route model<\/strong>: Theory that the first Paleoindians crossed to the Americas by following the southern coast of Beringia.<\/p>\n<p class=\"import-Normal\"><strong>Early Modern <\/strong><strong><em>Homo sapiens<\/em><\/strong><strong>, Early Anatomically Modern Human<\/strong>: Terms used to refer to transitional fossils between archaic and modern <em>Homo sapiens<\/em> that have a mosaic of traits. Humans like ourselves, who mostly lack archaic traits, are referred to as Late Modern <em>Homo sapiens<\/em> and simply Anatomically Modern Humans.<\/p>\n<p class=\"import-Normal\"><strong>Egalitarian<\/strong>: Human organization without strict ranks. Foraging societies tend to be more egalitarian than those based on other subsistence strategies.<\/p>\n<p class=\"import-Normal\"><strong>Foraging<\/strong>: Lifestyle consisting of frequent movement through the landscape and acquiring resources with minimal storage capacity.<\/p>\n<p class=\"import-Normal\"><strong>Generalist-specialist niche<\/strong>: The ability to survive in a variety of environments by developing local expertise. Evolution toward this niche may have been what allowed modern <em>Homo sapiens<\/em> to expand past the geographical range of other human species.<\/p>\n<p class=\"import-Normal\"><strong>Globalization<\/strong>: A recent increase in the interconnectedness and interdependence of people that is facilitated with long-distance networks.<\/p>\n<p class=\"import-Normal\"><strong>Globular<\/strong>: Having a rounded appearance. Increased globularity of the braincase is a trait of modern <em>Homo sapiens<\/em>.<\/p>\n<p class=\"import-Normal\"><strong>Gracile<\/strong>: Having a smooth and slender quality; the opposite of robust.<\/p>\n<p class=\"import-Normal\"><strong>Holocene<\/strong>: The epoch of the Cenozoic Era starting around 12,000 years ago and lasting arguably through the present.<\/p>\n<p class=\"import-Normal\"><strong>Ice-Free Corridor model<\/strong>: Theory that the first Native Americans crossed to the Americas through a passage between glaciers.<\/p>\n<p class=\"import-Normal\"><strong>Institutions<\/strong>: Long-lasting and influential cultural constructs. Examples include government, organized religion, academia, and the economy.<\/p>\n<p class=\"import-Normal\"><strong>Last Glacial Maximum<\/strong>: The time 23,000 years ago when the most recent ice age was the most intense.<\/p>\n<p class=\"import-Normal\"><strong>Later Stone Age<\/strong>: Time period following the Middle Stone Age with a diversification in tool types, starting around 50,000 years ago.<\/p>\n<p class=\"import-Normal\"><strong>Levant<\/strong>: The eastern coast of the Mediterranean. The site of early modern human expansion from Africa and later one of the centers of agriculture.<\/p>\n<p class=\"import-Normal\"><strong>Megafauna<\/strong>: Large ancient animals that may have been hunted to extinction by people around the world.<\/p>\n<p class=\"import-Normal\"><strong>Mental eminence<\/strong>: The chin on the mandible of modern <em>H. sapiens<\/em>. One of the defining traits of our species.<\/p>\n<p class=\"import-Normal\"><strong>Microlith<\/strong>: Small stone tool found in the Later Stone Age; also called a bladelet.<\/p>\n<p class=\"import-Normal\"><strong>Middle Stone Age<\/strong>: Time period known for Mousterian lithics that connects African archaic to modern <em>Homo sapiens<\/em>.<\/p>\n<p class=\"import-Normal\"><strong>Monumental architecture<\/strong>: Large and labor-intensive constructions that signify the power of the elite in a sedentary society. A common type is the pyramid, a raised crafted structure topped with a point or platform.<\/p>\n<p class=\"import-Normal\"><strong>Mosaic<\/strong>: Composed from a mix or composite of traits.<\/p>\n<p class=\"import-Normal\"><strong>Neolithic Revolution<\/strong>: Time of rapid change to human cultures due to the invention of agriculture, starting around 12,000 years ago.<\/p>\n<p class=\"import-Normal\"><strong>Ochre<\/strong>: Iron-based mineral pigment that can be a variety of yellows, reds, and browns. Used by modern human cultures worldwide since at least 80,000 years ago.<\/p>\n<p class=\"import-Normal\"><strong>Sahul<\/strong>: Ancient landmass connecting New Guinea and Australia.<\/p>\n<p class=\"import-Normal\"><strong>Sedentarism<\/strong>: Lifestyle based on having a stable home area; the opposite of nomadism.<\/p>\n<p class=\"import-Normal\"><strong>Southern Dispersal model<\/strong>: Theory that modern <em>H. sapiens<\/em> expanded from East Africa by crossing the Red Sea and following the coast east across Asia.<\/p>\n<p class=\"import-Normal\"><strong>Subsistence strategy<\/strong>: The method an organism uses to find nourishment and other resources.<\/p>\n<p class=\"import-Normal\"><strong>Sunda<\/strong>: Ancient Asian landmass that incorporated modern Southeast Asia.<\/p>\n<p class=\"import-Normal\"><strong>Supraorbital torus<\/strong>: The bony brow ridge across the top of the eye orbits on many hominin crania.<\/p>\n<p class=\"import-Normal\"><strong>Upper Paleolithic<\/strong>: Time period considered synonymous with the Later Stone Age.<\/p>\n<p class=\"import-Normal\"><strong>Urbanization<\/strong>: The increase of population density as people settled together in cities.<\/p>\n<p class=\"import-Normal\"><strong>Wallacea<\/strong>: Archipelago southeast of Sunda with different biodiversity than Asia.<\/p>\n<p class=\"import-Normal\"><strong>Younger Dryas<\/strong>: The rapid change in global climate\u2014notably a cooling of the Northern Hemisphere\u201413,000 years ago.<\/p>\n<h2 class=\"import-Normal\">About the Author<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\"><img loading=\"lazy\" decoding=\"async\" class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image14-3-1.jpg\" alt=\"A man with short, black hair stands outdoors with a setting sun behind him.\" width=\"282\" height=\"376\" \/><\/p>\n<h3 class=\"import-Normal\"><strong>Keith Chan, Ph.D.<\/strong><\/h3>\n<p class=\"import-Normal\">Grossmont-Cuyamaca Community College District and MiraCosta College, drkeithcchan@gmail.com, Dr. Keith Chan is an instructor of anthropology at Grossmont-Cuyamaca Community College District and MiraCosta College in San Diego County. He reached this step of his anthropological path after many memorable experiences across the country and the hemisphere. He earned a bachelor\u2019s degree in anthropology from the University of California, Berkeley, in 2001. As a graduate student at the University of Missouri, he traveled to Per\u00fa with teams of students to study skeletons in the archaeological record to understand the lives of ancient Andeans. He completed his dissertation and earned a Ph.D. in 2011. Inspired by many educators in his journey, Dr. Chan turned his career toward teaching anthropology and helping students understand and appreciate humanity.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration<\/h2>\n<h3 class=\"import-Normal\" style=\"text-indent: 0pt\"><strong>Websites<\/strong><\/h3>\n<p>First-person virtual tour of Lascaux cave with annotated cave art: Minist\u00e8re de la Culture and Mus\u00e9e d\u2019Arch\u00e9ologie Nationale. \u201c<a href=\"https:\/\/archeologie.culture.fr\/lascaux\/en\/visit-cave\" target=\"_blank\" rel=\"noopener\">Visit the cave<\/a>\u201d Lascaux website.<\/p>\n<p>Online anthropology magazine articles related to paleoanthropology and human evolution: SAPIENS. \u201c<a href=\"https:\/\/www.sapies.org\/category\/evolution\/\" target=\"_blank\" rel=\"noopener\">Evolution<\/a>.\u201d <em>SAPIENS<\/em> website.<\/p>\n<p>Various presentations of information about hominin evolution: Smithsonian Institution. \u201c<a href=\"https:\/\/humanorigins.si.edu\" target=\"_blank\" rel=\"noopener\">What does it mean to be human?<\/a>\u201d <em>Smithsonian National Museum of Natural History<\/em> website.<\/p>\n<p>Magazine-style articles on archaeology and paleoanthropology: ThoughtCo. \u201c<a href=\"https:\/\/www.thoughtco.com\/archaeology-4133504\" target=\"_blank\" rel=\"noopener\">Archaeology<\/a>.\u201d ThoughtCo. Website.<\/p>\n<p>Database of comparisons across hominins and primates: University of California, San Diego. \u201c<a href=\"https:\/\/carta.anthropogeny.org\/moca\/domains\" target=\"_blank\" rel=\"noopener\">MOCA Domains<\/a>.\u201d <em>Center for Academic Research &amp; Training in Anthropogeny<\/em> website.<\/p>\n<h3><strong>Books<\/strong><\/h3>\n<p>Engaging book that covers human-made changes to the environment with industrialization and globalization: Kolbert, Elizabeth. 2014. <em>The Sixth Extinction: An Unnatural History<\/em>. New York: Bloomsbury.<\/p>\n<p>Overview of what human life was like among the environmental shifts of the Ice Age: Woodward, Jamie. 2014. <em>The Ice Age: A Very Short Introduction<\/em>. Oxford: OUP Press.<\/p>\n<h3><strong>Articles<\/strong><\/h3>\n<p>Recent review paper about the current state of paleoanthropology research: Stringer, C. 2016. \u201c<a href=\"https:\/\/doi.org\/10.1098\/rstb.2015.0237\" target=\"_blank\" rel=\"noopener\">The Origin and Evolution of <em>Homo sapiens<\/em><\/a>.\u201d <em>Philosophical Transactions of the Royal Society B<\/em> 371 (1698).<\/p>\n<p>Overview of the history of American paleoanthropology and the many debates that have occurred over the years: Trinkaus, E. 2018. \u201cOne Hundred Years of Paleoanthropology: An American Perspective.\u201d <em>American Journal of Physical Anthropology<\/em> 165 (4): 638\u2013651.<\/p>\n<p>Amazing magazine article that synthesizes hominin evolution and why it is important to study this subject: Wheelwright, Jeff. 2015. \u201c<a href=\"https:\/\/discovermagazine.com\/2015\/may\/16-days-of-dysevolution\" target=\"_blank\" rel=\"noopener\">Days of Dysevolution<\/a>.\u201d <em>Discover<\/em> 36 (4): 33\u201339.<\/p>\n<p>Fascinating research on \u00d6tzi, a mummy from 5,000 years ago: Wierer, Ursula, Simona Arrighi, Stefano Bertola, G\u00fcnther Kaufmann, Benno Baumgarten, Annaluisa Pedrotti, Patrizia Pernter, and Jacques Pelegrin. 2018. \u201cThe Iceman\u2019s Lithic Toolkit: Raw Material, Technology, Typology and Use.\u201d <em>PLOS One<\/em> 13 (6): e0198292. https:\/\/doi.org\/10.1371\/journal.pone.0198292.<\/p>\n<h3><strong>Documentaries<\/strong><\/h3>\n<p>PBS NOVA series covering the expansion of modern <em>Homo sapiens<\/em> and interbreeding with archaic humans: Brown, Nicholas, dir. 2015. <em>First Peoples<\/em>. Edmonton: Wall to Wall Television. Amazon Prime Video.<\/p>\n<p>PBS NOVA special featuring the footprints found in White Sands National Park: Falk, Bella, dir. 2016. <em>Ice Age Footprints<\/em>. Boston: Windfall Films. https:\/\/www.pbs.org\/wgbh\/nova\/video\/ice-age-footprints\/.<\/p>\n<p>PBS NOVA special about how modern humans evolved adaptations to different environments. Shows how present-day people live around the world: Thompson, Niobe, dir. 2016. <em>Great Human Odyssey<\/em>. Edmonton: Clearwater Documentary. <a class=\"rId132\" href=\"https:\/\/www.pbs.org\/wgbh\/nova\/evolution\/great-human-odyssey.html\">https:\/\/www.pbs.org\/wgbh\/nova\/evolution\/great-human-odyssey.html<\/a>.<\/p>\n<\/div>\n<h2 class=\"__UNKNOWN__\">References<\/h2>\n<div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Araujo, Bernardo B. A., Luiz Gustavo R. Oliveira-Santos, Matheus S. Lima-Ribeiro, Jos\u00e9 Alexandre F. Diniz-Filho, and Fernando A. S. Fernandez. 2017. \u201cBigger Kill Than Chill: The Uneven Roles of Humans and Climate on Late Quaternary Megafaunal Extinctions.\u201d <em>Quaternary International<\/em> 431: 216\u2013222.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Armelagos, George J., Peter J. Brown, and Bethany Turner. 2005. \u201cEvolutionary, Historical, and Political Economic Perspectives on Health and Disease.\u201d <em>Social Science &amp; Medicine<\/em> 61 (4): 755\u2013765.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Armstrong, C. G., J. E. D. Miller, A. C. McAlvay, P. M. Ritchie, and D. Lepofsky. 2021. \u201cHistorical Indigenous Land-Use Explains Plant Functional Trait Diversity. <em>Ecology and Society<\/em> 26 (2): 6.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Bar-Yosef Mayer, Daniella E., Bernard Vandermeersch, and Ofer Bar-Yosef. 2009. \u201cShells and Ochre in Middle Paleolithic Qafzeh Cave, Israel: Indications for Modern Behavior.\u201d <em>Journal of Human Evolution<\/em> 56 (3): 307\u2013314.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Barbetti, M., and H. Allen. 1972. \u201cPrehistoric Man at Lake Mungo, Australia, by 32,000 Years Bp.\u201d <em>Nature<\/em> 240 (5375): 46\u201348.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Bennett, M. R., D. Bustos, J. S. Pigati, K. B. Springer, T. M. Urban, V. T. Holliday, Sally C. Reynolds, et al. (2021). \u201cEvidence of Humans in North America during the Last Glacial Maximum.\u201d <em>Science<\/em> 373 (6562): 1528\u20131531.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Bowler, J. M., Rhys Jones, Harry Allen, and A. G. Thorne. 1970. \u201cPleistocene Human Remains from Australia: A Living Site and Human Cremation from Lake Mungo, Western New South Wales.\u201d <em>World Archaeology<\/em> 2 (1): 39\u201360.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Brown, Peter. 1999. \u201cThe First Modern East Asians? Another Look at Upper Cave 101, Liujiang and Minatogawa 1.\u201d In <em>Interdisciplinary Perspectives on the Origins of the Japanese<\/em>, edited by K. Omoto, 105\u2013131. Kyoto: International Research Center for Japanese Studies.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Brown, Peter. 2000. \u201cAustralian Pleistocene Variation and the Sex of Lake Mungo 3.\u201d <em>Journal of Human Evolution<\/em> 38 (5): 743\u2013749.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Clarkson, Chris, Zenobia Jacobs, Ben Marwick, Richard Fullagar, Lynley Wallis, Mike Smith, Richard G. Roberts, et al. 2017. \u201cHuman Occupation of Northern Australia by 65,000 Years Ago.\u201d <em>Nature<\/em> 547 (7663): 306\u2013310.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Cohen, Mark Nathan. 1977. <em>The Food Crisis in Prehistory: Overpopulation and the Origins of Agriculture.<\/em> New Haven, CT: Yale University Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Cohen, Mark Nathan, and George J. Armelagos, eds. 1984.<em> Paleopathology at the Origins of Agriculture<\/em>. Orlando, FL: Academic Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Cohen, Mark Nathan, and Gillian M. M. Crane-Kramer, eds. 2007.<em> Ancient Health: Skeletal Indicators of Agricultural and Economic Intensification<\/em>. Gainesville, FL: University Press of Florida.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Copes-Gerbitz, K., S. Hagerman, and L. Daniels. 2021. \u201cSituating Indigenous Knowledge for Resilience in Fire-Dependent Social-Ecological Systems.\u201d <em>Ecology and Society<\/em> 26(4): 25. https:\/\/www.ecologyandsociety.org\/vol26\/iss4\/art25\/.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Coqueugniot, H\u00e9l\u00e8ne, Olivier Dutour, Baruch Arensburg, Henri Duday, Bernard Vandermeersch, and Anne-Marie Tillier. 2014. \u201cEarliest Cranio-Encephalic Trauma from the Levantine Middle Palaeolithic: 3-D Reappraisal of the Qafzeh 11 Skull, Consequences of Pediatric Brain Damage on Individual Life Condition and Social Care.\u201d <em>PLOS ONE<\/em> 9 (7): e102822.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Crittenden, Alyssa N., and Stephanie L. Schnorr. 2017. \u201cCurrent Views on Hunter\u2010Gatherer Nutrition and the Evolution of the Human Diet.\u201d <em>American Journal of Physical Anthropology<\/em> 162 (S63): 84\u2013109.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">d\u2019Errico, Francesco, Lucinda Backwell, Paola Villa, Ilaria Degano, Jeannette J. Lucejko, Marion K. Bamford, Thomas F. G. Higham, Maria Perla Colombini, and Peter B. Beaumont. 2012. \u201cEarly Evidence of San Material Culture Represented by Organic Artifacts from Border Cave, South Africa.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 109 (33): 13214\u201313219.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">d\u2019Errico, Francesco, Christopher Henshilwood, Marian Vanhaeren, and Karen Van Niekerk. 2005. \u201cNassarius Kraussianus Shell Beads from Blombos Cave: Evidence for Symbolic Behaviour in the Middle Stone Age.\u201d <em>Journal of Human Evolution<\/em> 48 (1): 3\u201324.<\/p>\n<p class=\"import-Normal\">Dannemann, Michael, and Fernando Racimo. 2018. \u201cSomething Old, Something Borrowed: Admixture and Adaptation in Human Evolution.\u201d <em>Current Opinion in Genetics &amp; Development<\/em> 53: 1\u20138.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Day, M. H. 1969. \u201cOmo Human Skeletal Remains.\u201d <em>Nature<\/em> 222: 1135\u20131138.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Dillehay, Tom D., Carlos Ocampo, Jos\u00e9 Saavedra, Andre Oliveira Sawakuchi, Rodrigo M. Vega, Mario Pino, Michael B. Collins, et al. 2015. \u201cNew Archaeological Evidence for an Early Human Presence at Monte Verde, Chile.\u201d <em>PLOS ONE<\/em> 10 (11): e0141923. doi:10.1371\/journal.pone.0141923.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Dow, Gregory K., Clyde G. Reed, and Nancy Olewiler. 2009. \u201cClimate Reversals and the Transition to Agriculture.\u201d <em>Journal of Economic Growth<\/em> 14 (1): 27\u201353.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Durband, Arthur C. 2014. \u201cBrief Communication: Artificial Cranial Modification in Kow Swamp and Cohuna.\u201d <em>American Journal of Physical Anthropology<\/em> 155 (1): 173\u2013178.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ember, Carol R. N.d. \u201cHunter-Gatherers.\u201d <em>Explaining Human Culture. Human Relations Area Files<\/em>. Accessed March 4, 2023. <a class=\"rId133\" href=\"https:\/\/hraf.yale.edu\/ehc\/summaries\/hunter-gatherers\">https:\/\/hraf.yale.edu\/ehc\/summaries\/hunter-gatherers<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Erlandson, Jon M., Todd J. Braje, Kristina M. Gill, and Michael H. Graham. 2015. \u201cEcology of the Kelp Highway: Did Marine Resources Facilitate Human Dispersal from Northeast Asia to the Americas?\u201d <em>The Journal of Island and Coastal Archaeology<\/em> 10 (3): 392\u2013411.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Fladmark, K. R. 1979. \u201cRoutes: Alternate Migration Corridors for Early Man in North America.\u201d <em>American Antiquity<\/em> 44 (1): 55\u201369.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Fletcher, M. S., T. Hall, and A. N. Alexandra. 2021. \u201cThe Loss of an Indigenous Constructed Landscape Following British Invasion of Australia: An Insight into the Deep Human Imprint on the Australian Landscape.\u201d <em>Ambio<\/em> 50(1): 138\u2013149.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Fu, Qiaomei, Mateja Hajdinjak, Oana Teodora Moldovan, Silviu Constantin, Swapan Mallick, Pontus Skoglund, Nick Patterson, et al. 2015. \u201cAn Early Modern Human from Romania with a Recent Neanderthal Ancestor.\u201d <em>Nature<\/em> 524 (7564): 216\u2013219.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Fuller, Dorian Q. 2010. \u201cAn Emerging Paradigm Shift in the Origins of Agriculture.\u201d <em>General Anthropology<\/em> 17 (2): 1, 8\u201311.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Gammage, B. 2008. \u201cPlain Facts: Tasmania under Aboriginal Management.\u201d <em>Landscape Research<\/em> 33 (2): 241\u2013254.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Germonpr\u00e9, Mietje, Martina L\u00e1zni\u010dkov\u00e1-Galetov\u00e1, and Mikhail V. Sablin. 2012. \u201cPalaeolithic Dog Skulls at the Gravettian P\u0159edmost\u00ed Site, the Czech Republic.\u201d <em>Journal of Archaeological Science<\/em> 39 (1): 184\u2013202.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Gr\u00f6ning, Flora, Jia Liu, Michael J. Fagan, and Paul O\u2019Higgins. 2011. \u201cWhy Do Humans Have Chins? Testing the Mechanical Significance of Modern Human Symphyseal Morphology with Finite Element Analysis.\u201d <em>American Journal of Physical Anthropology<\/em> 144 (4): 593\u2013606.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Harvati, Katerina. 2009. \u201cInto Eurasia: A Geometric Morphometric Reassessment of the Upper Cave (Zhoukoudian) Specimens.\u201d <em>Journal of Human Evolution<\/em> 57 (6): 751\u2013762.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Headland, Thomas N., Lawrence A. Reid, M. G. Bicchieri, Charles A. Bishop, Robert Blust, Nicholas E. Flanders, Peter M. Gardner, Karl L. Hutterer, Arkadiusz Marciniak, and Robert F. Schroeder. 1989. \u201cHunter-Gatherers and Their Neighbors from Prehistory to the Present.\u201d <em>Current Anthropology<\/em> 30 (1): 43\u201366.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Henshilwood, Christopher S., Francesco d\u2019Errico, Karen L. van Niekerk, Yvan Coquinot, Zenobia Jacobs, Stein-Erik Lauritzen, Michel Menu, and Renata Garc\u00eda-Moreno. 2011. \u201cA 100,000-Year-Old Ochre-Processing Workshop at Blombos Cave, South Africa.\u201d <em>Science<\/em> 334 (6053): 219\u2013222.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Hershkovitz, Israel, Gerhard W. Weber, Rolf Quam, Mathieu Duval, Rainer Gr\u00fcn, Leslie Kinsley, Avner Ayalon, et al. 2018. \u201cThe Earliest Modern Humans Outside Africa.\u201d <em>Science<\/em> 359 (6374): 456\u2013459.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Hublin, Jean-Jacques, Abdelouahed Ben-Ncer, Shara E. Bailey, Sarah E. Freidline, Simon Neubauer, Matthew M. Skinner, Inga Bergmann, et al. 2017. \u201cNew Fossils from Jebel Irhoud, Morocco, and the Pan-African Origin of <em>Homo sapiens<\/em>.\u201d <em>Nature<\/em> 546 (7657): 289\u2013292.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Lepofsky, D., N. F. Smith, N. Cardinal, J. Harper, M. Morris, M., Gitla (Elroy White), Randy Bouchard, et al. 2015. \u201cAncient Shellfish Mariculture on the Northwest Coast of North America.\u201d <em>American Antiquity<\/em> 80 (2): 236\u2013259.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Lieberman, Daniel E. 2015. \u201cHuman Locomotion and Heat Loss: An Evolutionary Perspective.\u201d <em>Comprehensive Physiology<\/em> 5 (1): 99\u2013117.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Lieberman, Daniel E., Brandeis M. McBratney, and Gail Krovitz. 2002. \u201cThe Evolution and Development of Cranial Form in <em>Homo sapiens<\/em>.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 99 (3): 1134\u20131139.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Lieberman, Daniel E., Osbjorn M. Pearson, and Kenneth M. Mowbray. 2000. \u201cBasicranial Influence on Overall Cranial Shape.\u201d <em>Journal of Human Evolution<\/em> 38 (2): 291\u2013315.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Liu, Wu, Mar\u00eda Martin\u00f3n-Torres, Yan-jun Cai, Song Xing, Hao-wen Tong, Shu-wen Pei, Mark Jan Sier, Xiao-hong Wu, R. Lawrence Edwards, and Hai Cheng. 2015. \u201cThe Earliest Unequivocally Modern Humans in Southern China.\u201d <em>Nature<\/em> 526 (7575): 696-699.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Lucas, Peter W. 2007. \u201cThe Evolution of the Hominin Diet from a Dental Functional Perspective.\u201d In <em>Evolution of the Human Diet: The Known, the Unknown, and the Unknowable<\/em>, edited by Peter S. Ungar, 31\u201338 Oxford, UK: Oxford University Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">McCarthy, Robert C., and Lynn Lucas. 2014. \u201cA Morphometric Reassessment of Bou-Vp-16\/1 from Herto, Ethiopia.\u201d <em>Journal of Human Evolution<\/em> 74: 114\u2013117.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">McDougall, Ian, Francis H. Brown, and John G. Fleagle. 2005. \u201cStratigraphic Placement and Age of Modern Humans from Kibish, Ethiopia.\u201d <em>Nature<\/em> 433 (7027): 733\u2013736.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">McNiven, I. J., J. Crouch, T. Richards, N. Dolby, and G. Jacobsen. 2012. \u201cDating Aboriginal Stone-Walled Fishtraps at Lake Condah, Southeast Australia.\u201d <em>Journal of Archaeological Science<\/em> 39 (2): 268\u2013286.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">McNiven, I., J. Crouch, T. Richards, K. Sniderman, N. Dolby, and G. Mirring. 2015. \u201cPhased Redevelopment of an Ancient Gunditjmara Fish Trap over the Past 800 Years: Muldoons Trap Complex, Lake Condah, Southwestern Victoria.\u201d <em>Australian Archaeology<\/em> 81 (1): 44\u201358.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Michel, V\u00e9ronique, H\u00e9l\u00e8ne Valladas, Guanjun Shen, Wei Wang, Jian-xin Zhao, Chuan-Chou Shen, Patricia Valensi, and Christopher J. Bae. 2016. \u201cThe Earliest Modern <em>Homo sapiens<\/em> in China?\u201d <em>Journal of Human Evolution<\/em> 101: 101\u2013104.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Miller, D. Shane, Vance T. Holliday, and Jordon Bright. 2013. \u201cClovis across the Continent.\u201d In <em>Paleoamerican Odyssey<\/em>, edited by Kelly E. Graf, Caroline V. Ketron, and Michael R. Waters, 207\u2013220. College Station: Texas A&amp;M University Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Neubauer, Simon, Jean-Jacques Hublin, and Philipp Gunz. 2018. \u201cThe Evolution of Modern Human Brain Shape.\u201d <em>Science Advances<\/em> 4 (1): eaao5961. https:\/\/doi.org\/10.1126\/sciadv.aao5961.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Pearson, Osbjorn M. 2000. \u201cPostcranial Remains and the Origin of Modern Humans.\u201d <em>Evolutionary Anthropology<\/em> 9: 229\u2013247.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Pearson, Osbjorn M. 2008. \u201cStatistical and Biological Definitions of \u2018Anatomically Modern\u2019 Humans: Suggestions for a Unified Approach to Modern Morphology.\u201d <em>Evolutionary Anthropology: Issues, News, and Reviews<\/em> 17 (1): 38\u201348.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Pietschnig, Jakob, Lars Penke, Jelte M. Wicherts, Michael Zeiler, and Martin Voracek. 2015. \u201cMeta-Analysis of Associations between Human Brain Volume and Intelligence Differences: How Strong Are They and What Do They Mean?\u201d <em>Neuroscience &amp; Biobehavioral Reviews<\/em> 57: 411\u2013432.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Posth, Cosimo, Nathan Nakatsuka, Iosif Lazaridis, Pontus Skoglund, Swapan Mallick, Thiseas C. Lamnidis, Nadin Rohland, et al. 2018. \u201cReconstructing the Deep Population History of Central and South America.\u201d <em>Cell<\/em> 175 (5): 1185\u20131197.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Potter, Ben A., James F. Baichtal, Alwynne B. Beaudoin, Lars Fehren-Schmitz, C. Vance Haynes, Vance T. Holliday, Charles E. Holmes, et al. 2018. \u201cCurrent Evidence Allows Multiple Models for the Peopling of the Americas.\u201d <em>Science Advances<\/em> 4 (8): eaat5473. <a class=\"rId134\" href=\"https:\/\/doi.org\/10.1126\/sciadv.aat5473\">https:\/\/doi.org\/10.1126\/sciadv.aat5473<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Reich, David, Richard E. Green, Martin Kircher, Johannes Krause, Nick Patterson, Eric Y. Durand, Bence Viola, et al. 2010. \u201cGenetic History of an Archaic Hominin Group from Denisova Cave in Siberia.\u201d <em>Nature<\/em> 468 (7327): 1053\u20131060.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Reich, David, Nick Patterson, Martin Kircher, Frederick Delfin, Madhusudan R. Nandineni, Irina Pugach, Albert Min-Shan Ko, et al. 2011. \u201cDenisova Admixture and the First Modern Human Dispersals into Southeast Asia and Oceania.\u201d <em>American Journal of Human Genetics<\/em> 89 (4): 516\u2013528.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Richter, Daniel, Rainer Gr\u00fcn, Renaud Joannes-Boyau, Teresa E. Steele, Fethi Amani, Mathieu Ru\u00e9, Paul Fernandes, et al. 2017. \u201cThe Age of the Hominin Fossils from Jebel Irhoud, Morocco, and the Origins of the Middle Stone Age.\u201d <em>Nature<\/em> 546 (7657): 293\u2013296.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Roberts, Patrick, and Brian A. Stewart. 2018. \u201cDefining the \u2018Generalist-Specialist\u2019 Niche for Pleistocene <em>Homo sapiens<\/em>.\u201d <em>Nature Human Behaviour<\/em> 2: 542\u2013550.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Rougier, Helene, \u015etefan Milota, Ricardo Rodrigo, Mircea Gherase, Lauren\u0163iu Sarcin\u01ce, Oana Moldovan, Jo\u00e3o Zilh\u00e3o, et al. 2007. \u201cPe\u015ftera Cu Oase 2 and the Cranial Morphology of Early Modern Europeans.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 104 (4): 1165\u20131170.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Sahlins, Marshall. 1968. \u201cNotes on the Original Affluent Society.\u201d In <em>Man the Hunter<\/em>, edited by R. B. Lee and I. DeVore, 85\u201389. New York: Aldine Publishing Company.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Sawyer, G. J., and Blaine Maley. 2005. \u201cNeanderthal Reconstructed.\u201d <em>The Anatomical Record (Part B: New Anat.)<\/em> 283 (1): 23\u201331.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scerri, Eleanor M. L., Mark G. Thomas, Andrea Manica, Philipp Gunz, Jay T. Stock, Chris Stringer, Matt Grove, et al. 2018. \u201cDid Our Species Evolve in Subdivided Populations Across Africa, and Why Does It Matter?\u201d <em>Trends in Ecology &amp; Evolution<\/em> 33 (8): 582\u2013594.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Shea, John J. 2011. \u201cRefuting a Myth about Human Origins.\u201d <em>American Scientist<\/em> 99 (2): 128\u2013135.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Shea, John J., and Ofer Bar-Yosef. 2005. \u201cWho Were the Skhul\/Qafzeh People? An Archaeological Perspective on Eurasia\u2019s Oldest Modern Humans.\u201d <em>Journal of the Israel Prehistoric Societ<\/em><em>y<\/em> 35: 451\u2013468.<\/p>\n<p class=\"import-Normal\">Slatkin, Montgomery, and Fernando Racimo. 2016. \u201cAncient DNA and Human History.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 113 (23): 6380\u20136387.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Smith, Fred H., James C. M. Ahern, Ivor Jankovi\u0107, and Ivor Karavani\u0107. 2017. \u201cThe Assimilation Model of Modern Human Origins in Light of Current Genetic and Genomic Knowledge.\u201d <em>Quaternary International<\/em> 450: 126\u2013136.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Smith, Michael. 2009. \u201cV. Gordon Childe and the Urban Revolution: A Historical Perspective on a Revolution in Urban Studies.\u201d <em>Town Planning Review<\/em> 80 (1): 3\u201329.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Stock, Jay T. 2008. \u201cAre Humans Still Evolving?\u201d <em>EMBO Reports<\/em> 9 (Suppl 1): S51\u2013S54.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Swisher, Mark E., Dennis L. Jenkins, Lionel E. Jackson Jr., and Fred M. Phillips. 2013. \u201cA Reassessment of the Role of the Canadian Ice-Free Corridor in Light of New Geological Evidence.\u201d Poster Symposium 5B: Geology, Geochronology and Paleoenvironments of the First Americans at the Paleoamerican Odyssey Conference, Santa Fe, New Mexico, October 16\u201319.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Thorne, A. G., and P. G. Macumber. 1972. \u201cDiscoveries of Late Pleistocene Man at Kow Swamp, Australia.\u201d <em>Nature<\/em> 238 (5363): 316\u2013319.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Trinkaus, Erik, \u015etefan Milota, Ricardo Rodrigo, Gherase Mircea, and Oana Moldovan. 2003a. \u201cEarly Modern Human Cranial Remains from the Pe\u015ftera Cu Oase, Romania.\u201d <em>Journal of Human Evolution<\/em> 45 (3): 245\u2013253.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Trinkaus, Erik, Oana Moldovan, Adrian B\u00eelg\u0103r, Lauren\u0163iu Sarcina, Sheela Athreya, Shara E Bailey, Ricardo Rodrigo, Gherase Mircea, Thomas Higham, and Christopher Bronk Ramsey. 2003b. \u201cAn Early Modern Human from the Pe\u015ftera Cu Oase, Romania.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 100 (20): 11231\u201311236.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Velem\u00ednsk\u00e1, J., J. Br\u016fzek, P. Velem\u00ednsk\u00fd, L. Bigoni, A. Sefc\u00e1kov\u00e1, and S. Katina. 2008. \u201cVariability of the Upper-Palaeolithic Skulls from Predmost\u00ed Near Prerov (Czech Republic): Craniometric Comparison with Recent Human Standards.\u201d <em>Homo<\/em> 59 (1): 1\u201326.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Vidal, C\u00e9line M., Christine S. Lane, Asfawossen Asrat, Dan N. Barfod, Darren F. Mark, Emma L. Tomlinson, Ambdemichael Zafu Tadesse, et al. (2022). \u201cAge of the Oldest Known <em>Homo sapiens<\/em> from Eastern Africa. <em>Nature<\/em> 601 (7894): 579\u2013583.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Villa, Paola, Sylvain Soriano, Tsenka Tsanova, Ilaria Degano, Thomas F. G. Higham, Francesco d\u2019Errico, Lucinda Backwell, Jeannette J. Lucejko, Maria Perla Colombini, and Peter B. Beaumont. 2012. \u201cBorder Cave and the Beginning of the Later Stone Age in South Africa.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 109 (33): 13208\u201313213.<\/p>\n<p class=\"import-Normal\">Wall, Jeffrey D., and Deborah Yoshihara Caldeira Brandt. 2016. \u201cArchaic Admixture in Human History.\u201d <em>Current Opinion in Genetics &amp; Development<\/em> 41: 93\u201397.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">White, Tim D., Berhane Asfaw, David DeGusta, Henry Gilbert, Gary D. Richards, Gen Suwa, and F. Clark Howell. 2003. \u201cPleistocene <em>Homo sapiens<\/em> from Middle Awash, Ethiopia.\u201d <em>Nature<\/em> 423 (6941): 742\u2013747.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Woo, Ju-Kang. 1959. \u201cHuman Fossils Found in Liukiang, Kwangsi, China.\u201d <em>Vertebrata PalAsiatica<\/em> 3 (3): 109\u2013118.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Wu, XiuJie, Wu Liu, Wei Dong, JieMin Que, and YanFang Wang. 2008. \u201cThe Brain Morphology of Homo Liujiang Cranium Fossil by Three-Dimensional Computed Tomography.\u201d <em>Chinese Science Bulletin<\/em> 53 (16): 2513\u20132519.<\/p>\n<h2 class=\"import-Normal\">Acknowledgments<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">I could not have undertaken this project without the help of many who got me to where I am today. I extend sincere thank yous to the many colleagues and former students who have inspired me to keep learning and talking about anthropology. Thank you also to all who are involved in this textbook project. The anonymous reviewers truly sparked improvements to the chapter. Lastly, the staff of Starbucks #5772 also contributed immensely to this text.<\/p>\n<\/div>\n<div class=\"glossary\"><span class=\"screen-reader-text\" id=\"definition\">definition<\/span><template id=\"term_382_2708\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_2708\"><div tabindex=\"-1\"><\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1406\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1406\"><div tabindex=\"-1\"><p>Slender, less rugged, or pronounced features.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1758\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1758\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\">Jonathan Marks, Ph.D., University of North Carolina at Charlotte<\/p>\n<p class=\"import-Normal\">Adam P. Johnson, M.A., University of North Carolina at Charlotte\/University of Texas at San Antonio<\/p>\n<p class=\"import-Normal\"><em>This chapter is an adaptation of \"<\/em><a class=\"rId9\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__\/\"><em>Chapter 2: Evolution<\/em><\/a><em>\u201d by Jonathan Marks. In <\/em><a class=\"rId10\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, first edition<\/em><\/a><em>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under <\/em><a class=\"rId11\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\"><em>CC BY-NC 4.0<\/em><\/a><em>. <\/em><\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\"><span style=\"color: #ffffff\">Learning Objectives<\/span><\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li>Explain the relationship among genes, bodies, and organismal change.<\/li>\n<li>Discuss the shortcomings of simplistic understandings of genetics.<\/li>\n<li>Describe what is meant by the \"biopolitics of heredity.\"<\/li>\n<li>Discuss issues caused by misuse of ideas about adaptations and natural selection.<\/li>\n<li>Examine and correct myths about evolution.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p class=\"import-Normal\">The Human Genome Project, an international initiative launched in 1990, sought to identify the entire genetic makeup of our species. For many scientists, it meant trying to understand the genetic underpinnings of what made humans uniquely human. James Watson, a codiscoverer of the helical shape of DNA, wrote that \u201cwhen finally interpreted, the genetic messages encoded within our DNA molecules will provide the ultimate answers to the chemical underpinnings of human existence\u201d (Watson 1990, 248). The underlying message is that what makes humans unique can be found in our <strong>genes<\/strong>. The Human Genome Project hoped to find the core of who we are and where we come from.<\/p>\n<p class=\"import-Normal\">Despite its lofty goal, the Human Genome Project\u2014even after publishing the entire human genome in January 2022\u2014could not fully account for the many factors that contribute to what it is to be human. Richard Lewontin, Steven Rose, and Leon Kamin (2017) argue that genetic determinism of the sort assumed by the Human Genome Project neglects other essential dimensions that contribute to the development and evolution of human bodies, not to mention the role that culture plays. They use an apt metaphor of a cake to illustrate the incompleteness of reductive models. Consider the flavor of a cake and think of the ingredients listed in the recipe. The recipe includes ingredients such as flour, sugar, shortening, vanilla extract, eggs, and milk. Does raw flour taste like cake? Does sugar, vanilla extract, or any of the other ingredients taste like cake? They do not, and knowing the individual flavors of each ingredient does not tell us much about what cake tastes like. Even mixing all of the ingredients in the correct proportions does not get us cake. Instead, external factors such as baking at the right temperature, for the right amount of time, and even the particularities of our evolved sense of taste and smell are all necessary components of experiencing the cake.<\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ff00ff\">Lewontin, Rose, and Kamin (2017) argue that the same is true for humans and other organisms.<\/span><\/p>\n<p class=\"import-Normal\">Knowing everything about cake ingredients does not allow us to fully know cake. Equally so, knowing everything about the genes found in our DNA does not allow us to fully know humans. Different, interacting levels are implicated in the development and evolution of all organisms, including humans. Genes, the structure of chromosomes, developmental processes, epigenetic tags, environmental factors, and still-other components all play key roles such that genetically reductive models of human development and evolution are woefully inadequate.<\/p>\n<p class=\"import-Normal\">The complex interactions across many levels\u2014genetic, developmental, and environmental\u2014explain why we still do not know how our one-dimensional DNA nucleotide sequence results in a four-dimensional organism. This was the unfulfilled promise of the inception of the Human Genome Project in the 1980s and 1990s: the project produced the complete DNA sequence of a human cell in the hopes that it would reveal how human bodies are built and how to cure them when they are built poorly. Yet, that information has remained elusive. Presumably, the knowledge of how organisms are produced from DNA sequences will one day permit us to reconcile the discrepancies between patterns in anatomical evolution and molecular evolution.<\/p>\n<p class=\"import-Normal\">In this chapter, we will consider multilevel evolution and explore evolution as a complex interaction between genetic and epigenetic factors as well as the environments in which organisms live. Next, we will examine the biopolitical nature of human evolution. We will then investigate problems that arise from attributing all traits to an adaptive function. Finally, we will address common misconceptions about evolution. The goal of this chapter is to provide you with the necessary toolkit for understanding the molecular, anatomical, and political dimensions of evolution.<\/p>\n<h2 class=\"import-Normal\">Evolution Happens at Multiple Levels<\/h2>\n<p class=\"import-Normal\">Following Richard Dawkins\u2019s publication of <em>The Selfish Gene <\/em>in 1976, the scientific imagination was captured by the potential of genomics to reveal how genes are copied by Darwinian selection. Dawkins argues that the genes in individuals that contribute to greater reproductive success are the units of selection. His conception of evolution at the molecular level undercuts the complex interactions between organisms and their environments, which are not expressed genomically but are nevertheless key drivers in evolution.<\/p>\n<p class=\"import-Normal\">By the 1980s, the acknowledgment among most biologists that even though genes construct bodies, genes and bodies evolve at different rates and with distinct patterns. This realization led to a renewed focus on how bodies change. The Evolutionary Synthesis of the 1930s\u20131970s had reduced organisms to their <strong>genotypes<\/strong> and species to their <strong>gene pools<\/strong>, which provided valuable insights about the processes of biological change, but it was only a first approximation. Animals are in fact reactive and adaptable beings, not passive and inert genotypes. Species are clusters of socially interacting and reproductively compatible organisms.<\/p>\n<figure style=\"width: 291px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/06\/image8-5.png\" alt=\"An asteroid hits the ocean. Pterodactyls fly among clouds in the foreground.\" width=\"291\" height=\"233\" \/><figcaption class=\"wp-caption-text\">Figure 17.1: A painting by Donald E. Davis representing the Chicxulub asteroid impact off the Yucatan Peninsula that contributed to the mass extinction that included the dinosaurs about 65 million years ago. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Chicxulub_impact_-_artist_impression.jpg\">Chicxulub impact - artist impression<\/a> by Donald E. Davis, <a href=\"https:\/\/www.nasa.gov\/\">NASA<\/a>, is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Once we accept that evolutionary change is fundamentally genetic change, we can ask: How do bodies function and evolve? How do groups of animals come to see one another as potential mates or competitors for mates, as opposed to just other creatures in the environment? Are there evolutionary processes that are not explicable by population genetics? These questions\u2014which lead us beyond reductive assumptions\u2014were raised in the 1980s by Stephen Jay Gould, the leading evolutionary biologist of the late 20th century (see: Gould 2003; 1996).<\/p>\n<p class=\"import-Normal\">Gould spearheaded a movement to identify and examine higher-order processes and features of evolution that were not adequately explained by population genetics. For example, <strong>extinction<\/strong>, which was such a problem for biologists of the 1600s, could now be seen as playing a more complex role in the history of life than population genetics had been able to model. Gould recognized that there are two kinds of extinctions, each with different consequences: background extinctions and mass extinctions. Background extinctions are those that reflect the balance of nature, because in a competitive Darwinian world, some things go extinct and other things take their place. Ecologically, your species may be adapted to its niche, but if another species comes along that\u2019s better adapted to the same niche, eventually your species will go extinct. It sucks, but it is the way of all life: you come into existence, you endure, and you pass out of existence. But mass extinctions are quite different. They reflect not so much the balance of nature as the wholesale disruption of nature: many species from many different lineages dying off at roughly the same time\u2014presumably as the result of some kind of rare ecological disaster. The situation may not be survival of the fittest as much as survival of the luckiest. The result, then, would be an ecological scramble among the survivors. Having made it through the worst, the survivors could now simply divide up the new ecosystem amongst themselves, since their competitors were gone. Something like this may well have happened about 65 million years ago, when a huge asteroid hit the Yucatan Peninsula, which mammals survived but dinosaurs did not (Figure 17.1). Something like this may be happening now, due to human expansion and environmental degradation. Note, though, that there is only a limited descriptive role here for population genetics: the phenomena we are describing are about organisms and species in ecosystems.<\/p>\n<p class=\"import-Normal\">Another question involved the disconnect between properties of <em>species<\/em> and the properties of <em>gene pools<\/em>. For example, there are upwards of 15 species of gibbons but only two species of chimpanzees. Why? There are upwards of 20 species of guenons but fewer than ten of baboons. Why? Are there genes for that? It seems unlikely. Gould suggested that species, as units of nature, might have properties that are not reducible to the genes in their cells. For example, rates of speciation and extinction might be properties of their ecologies and histories rather than their genes. Thus, relationships between environmental contexts and variability within a species result in degrees of resistance to extinction and affect the frequency and rates at which clades diversify (Lloyd and Gould 1993). Consistent biases of speciation rates might well produce patterns of macroevolutionary diversity that are difficult to explain genetically and better understood ecologically. Gould called such biases in speciation rates <strong>species selection<\/strong>\u2014a higher-order process that invokes competition between species, in addition to the classic Darwinian competition between individuals.<\/p>\n<p class=\"import-Normal\">One of Gould\u2019s most important studies involved the very nature of species. In the classical view, a species is continually adapting to its environment until it changes so much that it is a different species than it was at the beginning of this sentence (Eldredge and Gould 1972). That implies that the species is a fundamentally unstable entity through time, continuously changing to fit in. But suppose, argued Gould along with paleontologist Niles Eldredge, a species is more stable through time and only really adapts during periods of ecological instability and change. Then we might expect to find in the fossil record long equilibrium periods\u2014a few million years or so\u2014in which species don\u2019t seem to change much, punctuated by relatively brief periods in which they change a bit and then stabilize again as new species. They called this idea <strong>punctuated equilibria<\/strong>. The idea helps to explain certain features of the fossil record, notably the existence of small anatomical \u201cgaps\u201d between closely related fossil forms (Figure 17.2). Its significance lies in the fact that although it incorporates genetics, punctuated equilibria is not really a theory of genetics but one of types bodies in deep time.<\/p>\n<p class=\"import-Normal\">Punctuated equilibria is seen across taxa, with long periods in the fossil record representing little phenotypic change. These periods of stability are disrupted by shorter periods of rapid <strong>adaptation<\/strong>, the process through which populations of organisms become suited to living in their environments. Phenotypic changes are often coupled with drastic climatic or ecological changes that affect the milieu in which organisms live. For example, throughout much of hominin evolutionary history, brain size was closely associated with body size and thus remained mostly stable. However, changes occurred in average hominin brain size at around 100 thousand years ago, 1 million years ago, and 1.8 million years ago. Several hypotheses have been put forth to explain these changes, including unpredictability in climate and environment (Potts 1998), social development (Barton 1996), and the evolution of language (Deacon 1998). Evidence from the fossil record, paleoclimate models, and comparative anatomy suggests that the changes observed in hominin lineage result from biocultural processes\u2014that is, the coalescence of environmental and cultural factors that selected for larger brains (Marks 2015; Shultz, Nelson, and Dunbar 2012).<\/p>\n<figure style=\"width: 461px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image5-8.png\" alt=\"Two graphs contrast phyletic gradualism and punctuated equilibria.\" width=\"461\" height=\"222\" \/><figcaption class=\"wp-caption-text\">Figure 17.2: Different ways of conceptualizing the evolutionary relationship between an earlier and a later species. With phyletic gradualism, species are envisioned transforming continually in a direct line over time. With punctuated equilibria species branch off at particular points over time.\u00a0 Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__\/\">Phyletic gradualism vs. punctuated equilibria (Figure 2.12)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">In response to the call for a theory of the evolution of form, the field of <strong>evo-devo<\/strong>\u2014the intersection of evolutionary and developmental biology\u2014arose. The central focus here is on how changes in form and shape arise. An embryo matures by the stimulation of certain cells to divide, forming growth fields. The interactions and relationships among these growth fields generate the structures of the body. The <strong>hox genes<\/strong> that regulate these growth fields turn out to be highly conserved across the animal kingdom. This is because they repeatedly turn on and off the most basic genes guiding the animal\u2019s development, and thus any changes to them would be catastrophic. Indeed, these genes were first identified by manipulating them in fruit flies, such that one could produce a bizarre mutant fruit fly that grew a pair of legs where its antennae were supposed to be (Kaufman, Seeger, and Olsen 1990).<\/p>\n<p class=\"import-Normal\">Certain genetic changes can alter the fates of cells and the body parts, while other genetic changes can simply affect the rates at which neighboring groups of cells grow and divide, thus producing physical bumps or dents in the developing body. The result of altering the relationships among these fields of cellular proliferation in the growing embryo is <strong>allometry<\/strong>, or the differential growth of body parts. As an animal gets larger\u2014either over the course of its life or over the course of macroevolution\u2014it often has to change shape in order to live at a different size. Many important physiological functions depend on properties of geometric area: the strength of a bone, for example, is proportional to its cross-sectional area. But area is a two-dimensional quality, while growing takes place in three dimensions\u2014as an increase in mass or volume. As an animal expands, its bones necessarily weaken, because volume expands faster than area does. Consequently a bigger animal has more stress on its bones than a smaller animal does and must evolve bones even thicker than they would be by simply scaling the animal up proportionally. In other words, if you expand a mouse to the size of an elephant, it will nevertheless still have much thinner bones than the elephant does. But those giant mouse bones will unfortunately not be adequate to the task. Thus, a giant mouse would have to change aspects of its form to maintain function at a larger size (see Figure 17.3).<\/p>\n<p class=\"import-Normal\"><img class=\"aligncenter\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image6-6.png\" alt=\"Side-view of a mouse skeleton.\" width=\"515\" height=\"252\" \/><\/p>\n<figure style=\"width: 453px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image7-9.png\" alt=\"Side-view of an elephant skeleton.\" width=\"453\" height=\"326\" \/><figcaption class=\"wp-caption-text\">Figure 17.3: Mouse (top) and elephant (bottom) skeletons. Notice the elephant\u2019s bones are more robust when the two animals are the same size. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__\/\">Mouse and elephant skeletons (Figure 2.13)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Physiologically, we would like to know how the body \u201cknows\u201d when to turn on and off the genes that regulate growth to produce a normal animal. Evolutionarily, we would like to know how the body \u201clearns\u201d to alter the genetic on\/off switch (or the genetic \u201cslow down\/speed up\u201d switch) to produce an animal that looks different. Moreover, since organisms differ from one another, we would like to know how the developing body distinguishes a range of normal variation from abnormal variation. And, finally, how does abnormal variation eventually become normal in a descendant species?<\/p>\n<p class=\"import-Normal\">Taking up these questions, Gould invoked the work of a British geneticist named Conrad H. Waddington, who thought about genetics in less reductive ways than his colleagues. Rather than isolate specific DNA sites to analyze their function, Waddington instead studied the inheritance of an organism\u2019s reactivity\u2014its ability to adapt to the circumstances of its life. In a famous experiment, he grew fruit fly eggs in an atmosphere containing ether. Most died, but a few survived somehow by developing a weird physical feature: a second thorax with a second pair of wings. Waddington bred these flies and soon developed a stable line of flies who would reliably develop a second thorax when grown in ether. Then he began to lower the concentration of ether, while continuing to selectively breed the flies that developed the strange appearance. Eventually he had a line of flies that would stably develop the \u201cbithorax\u201d <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1238\"><strong>phenotype<\/strong><\/a>\u2013the suite of traits of an organism\u2013even when there was no ether; it had become the \u201cnew normal.\u201d The flies had genetically assimilated the bithorax condition.<\/p>\n<p class=\"import-Normal\">Waddington was thus able to mimic the <strong>inheritance of acquired characteristics<\/strong>: what had been a trait stimulated by ether a few generations ago was now a normal part of the development of the descendants. Waddington recognized that while he had performed a selection experiment on genetic variants, he had not selected for particular traits. Rather, he helped produce the physiological tendency to develop particular traits when appropriately stimulated. He called that tendency <strong>plasticity<\/strong> and its converse, the tendency to stay the same even under weird environmental circumstances, <strong>canalization.<\/strong> Waddington had initially selected for plasticity, the tendency to develop the bithorax phenotype under weird conditions, and then, later, for canalization, the developmental normalization of that weird physical trait. Although Waddington had high stature in the community of geneticists, evolutionary biologists of the 1950s and 1960s regarded him with suspicion because he was not working within the standard mindset of reductionism, which saw evolution as the spread of genetic variants that coded for favorable traits. Both Waddington and Gould resisted contemporary intellectual paradigms that favored reductive accounts of evolutionary processes. They conceived of evolution as an emergent process in which many external factors (e.g. climate, environment, predation) and internal factors (e.g., genotypes, plasticity, canalization) coalesce to produce the evolutionary trends that we observe in the fossil record and our genome.<\/p>\n<p class=\"import-Normal\">While Gould and Waddington both looked beyond the genome to understand evolution, the Human Genome Project\u2014an international project with the goal of identifying each base pair in the human genome in the 1990s\u2014generated a great deal of public interest in analyzing the human DNA sequence from the standpoint of medical genetics. Some of the rhetoric aimed to sell the public on investing a lot of money and resources in sequencing the human genome in order to show the genetic basis of heritable traits, cure genetic diseases, and learn what it means ultimately to be biologically human. However, the Human Genome Project was not actually able to answer those questions through the use of genetics alone, and thus a broader, more holistic account was required.<\/p>\n<p class=\"import-Normal\">This holistic account came from decades of research in human biology and anthropology, which understood the human body as highly adaptable, dynamic, and emergent. For example, in the early 20th century, anthropologist Franz Boas measured the skulls of immigrants to the U.S., revealing that environmental, not merely genetic, factors affected skull shape. The growing human body adjusts itself to the conditions of life, such as diet, sunshine, high altitude, hard labor, population density, how babies are carried\u2014any and all of which can have subtle but consistent effects upon its development. There can thus be no normal human form, only a context-specific range of human forms.<\/p>\n<p class=\"import-Normal\">However, what the human biologists called human adaptability, evolutionary biologists called developmental plasticity, and evidence quickly began to mount for its cause being <strong>epigenetic <\/strong>modifications to DNA. Epigenetic modifications are changes to how genes are used by the body (as opposed to changes in the DNA sequences; see Chapter 3). Scientific interest shifted from the focus of the Human Genome Project to the ways that bodies are made by evolutionary-developmental processes, including epigenetics. What is meant by \u201cepigenetic modification\u201d? Evolution is about how descendants diverge from their ancestors. Inheritance from parent to offspring is still critical to this process, which occurs through genetic recombination: the pairing of homologous chromosomes and sharing of genetic material during meiosis (see Chapter 3). However, in the 21st century, the link between evolution and inheritance has broadened with a clearer understanding of how environmental and developmental factors shape bodies and the expression of genes, including epigenetic inheritance patterns. While offspring inherit their genes through random assortment during meiosis, environmental factors also shape how genes are used. When these epigenetic modifications occur in germ cells, they can be passed onto offspring. In these cases, there is no change in the DNA sequence but rather in how genes are used by the body due to DNA methylation and the structure of chromosomes due to histone acetylation (see Chapter 3).<\/p>\n<p class=\"import-Normal\">In addition, we now recognize that evolution is affected by two other forms of intergenerational transmission and inheritance (in addition to genetics and epigenetics). These forms include behavioral variation and culture. That is, behavioral information can be transmitted horizontally (intragenerationally), permitting more rapid ways for organisms to adjust to the environment. And, then there is the fourth mode of transmission: the cultural or symbolic mode. <span style=\"background-color: #ffff00\">Humans are the only species<\/span> that horizontally transmits an arbitrary set of rules to govern communication, social interaction, and thought. This shared information is symbolic and has resulted in what we recognize as \u201cculture\u201d: locally emergent worlds of names, words, pictures, classifications, revered pasts, possible futures, spirits, dead ancestors, unborn descendants, in-laws, politeness, taboo, justice, beauty, and story, all accompanied by practices and a material world of tools.<\/p>\n<p class=\"import-Normal\">Consequently our contemporary ideas about evolution see the evolutionary processes as hierarchically organized and not restricted to the differential transmission of DNA sequences into the next generation. While that is indeed a significant part of evolution, the organism and species are nevertheless crucial to understanding how those DNA sequences get transmitted. Further, the transmission of epigenetic, behavioral, and symbolic information play a complex role in perpetuating our genes, bodies, and species. In the case of human evolution, one can readily see that symbolic information and cultural adaptation are far more central to our lives and our survival today than DNA and genetic adaptation. It is thus misleading to think of humans passively occupying an environmental niche. Rather, humans are actively engaged in constructing our own niches, as well as adapting to them and using them to adapt. The complex interplay between a species and its active engagement in creating its own ecology is known as <strong>niche construction<\/strong>. If we understand <strong>natural selection<\/strong>\u2013the process by which populations adapt to their specific environments\u2013as the effects that environmental context has on the reproductive success of organisms, then niche construction is the process through which organisms shape their own selective pressures.<\/p>\n<h2 class=\"import-Normal\">The Biopolitics of Heredity<\/h2>\n<p class=\"import-Normal\">\u201cScience isn\u2019t political\u201d is a sentiment that you have likely heard before. Science is supposed to be about facts and objectivity. It exists, or at least ought to, outside of petty human concerns. However, the sorts of questions we ask as scientists, the problems we choose to study, the categories and concepts we use, who gets to do science, and whose work gets cited are all shaped by culture. Doing science is a political act. This fact is markedly true for human evolution. While it is easier to create intellectual distance between us and fruit flies and viruses, there is no distance when we are studying ourselves. The hardest lesson to learn about human evolution is that it is intensely political. Indeed, to see it from the opposite side, as it were, the history of creationism\u2014the belief that the universe was divinely created around 6,000 years ago\u2014is essentially a history of legal decisions. For instance, in <em>Tennessee v. John T. Scopes<\/em> (1925), a schoolteacher was prosecuted for violating a law in Tennessee that prohibited the teaching of human evolution in public schools, where teachers were required by law to teach creationism.<\/p>\n<p class=\"import-Normal\">More recently, legal decisions aimed at legislating science education have shaped how students are exposed to evolutionary theory. For instance, <em>McLean v. Arkansas<\/em> (1982) dispatched \u201cscientific creationism\u201d by arguing that the imposition of balanced teaching of evolution and creationism in science classes violates the Establishment Clause, separating church and state. Additionally, <em>Kitzmiller v. Dover (Pennsylvania) Area School District<\/em> (2005) dispatched the teaching of \u201cintelligent design\u201d in public school classrooms as it was deemed to not be science. In some cases, people see unbiblical things in evolution, although most Christian theologians are easily able to reconcile science to the Bible. In other cases, people see immoral things in evolution, although there is morality and immorality everywhere. And some people see evolution as an aspect of alt-religion, usurping the authority of science in schools to teach the rejection of the Christian faith, which would be unconstitutional due to the protected separation of church and state.<\/p>\n<p class=\"import-Normal\">Clearly, the position that politics has nothing to do with science is untenable. But is the politics in evolution an aberration or is it somehow embedded in science? In the early 20th century, scientists commonly promoted the view that science and politics were separate: science was seen as a pure activity, only rarely corrupted by politics. And yet as early as World War I, the politics of nationalism made a hero of the German chemist Fritz Haber for inventing poison gas. And during World War II, both German doctors and American physicists, recruited to the war effort, helped to end many civilian lives. Therefore, we can think of the apolitical scientist as a self-serving myth that functions to absolve scientists of responsibility for their politics. The history of science shows how every generation of scientists has used evolutionary theory to rationalize political and moral positions. In the very first generation of evolutionary science, Darwin\u2019s <em>Origin of Species<\/em> (1859) is today far more readable than his <em>Descent of Man<\/em> (1871). The reason is that Darwin consciously purged <em>The Origin of Species<\/em> of any discussion of people. And when he finally got around to talking about people, in <em>The Descent of Man<\/em>, he simply imbued them with the quaint Victorian prejudices of his age, and the result makes you cringe every few pages. There is plenty of politics in there\u2014sexism, racism, and colonialism\u2014because <em>you cannot talk about people apolitically<\/em>.<\/p>\n<p class=\"import-Normal\">One immediate faddish deduction from Darwinism, popularized by Herbert Spencer (1864) as \u201csurvival of the fittest,\u201d held that unfettered competition led to advancement in nature and to human history. Since the poor were purported losers in that struggle, anything that made their lives easier would go against the natural order. This position later came to be known ironically as \u201cSocial Darwinism.\u201d Spencer was challenged by fellow Darwinian Thomas Huxley (1863), who agreed that struggle was the law of the jungle but observed that we don\u2019t live in jungles anymore. The obligation to make lives better for others is a moral, not a natural, fact. We simultaneously inhabit a natural universe of descent from apes and a moral universe of injustice and inequality, and science is not well served by ignoring the latter.<\/p>\n<p class=\"import-Normal\">Concurrently, the German biologist Ernst Haeckel\u2019s 1868 popularization of Darwinism was translated into English a few years later as <em>The History of Creation<\/em>. As we saw earlier, Haeckel was determined to convince his readers that they were descended from apes, even in the absence of fossil evidence attesting to it. When he made non-Europeans into the missing links that connected his readers to the apes, and depicted them as ugly caricatures, he knew precisely what he was doing. Indeed, even when the degrading racial drawings were deleted from the English translation of his book, the text nevertheless made his arguments quite clear. And a generation later, when the Americans had not yet entered the Great War in 1916, a biologist named Vernon Kellogg visited the German High Command as a neutral observer and found that the officers knew a lot about evolutionary biology, which they had gotten from Haeckel and which rationalized their military aggressions. Kellogg went home and wrote a bestseller about it, called <em>Headquarters Nights<\/em> (1917). World War I would have been fought with or without evolutionary theory, but as a source of scientific authority, evolution\u2014even if a perversion of the Darwinian theory\u2014had very quickly attained global geopolitical relevance.<\/p>\n<p class=\"import-Normal\">Oftentimes, politics in evolutionary science is subtle, due to the pervasive belief in the advancement of science. We recognize the biases of our academic ancestors and modify our scientific stories accordingly. But we can never be free of our own cultural biases, which are invisible to us, as much as our predecessors\u2019 biases were invisible to them. In some cases, the most important cultural issues resurface in different guises each generation, like scientific racism. <strong>Scientific racism<\/strong> is the recruitment of science for the evil political ends of racism, and it has proved remarkably impervious to evolution. Before Darwin, there was creationist scientific racism, and after Darwin, there was evolutionist scientific racism. And there is still scientific racism today, self-justified by recourse to evolution, which means that scientists have to be politically astute and sensitive to the uses of their work to counter these social tendencies.<\/p>\n<p class=\"import-Normal\">Consider this: Are you just your ancestry, or can you transcend it? If that sounds like a weird question, it was actually quite important to a turn-of-the-20th-century European society in which an old hereditary aristocracy was under increasing threat from a rising middle class. And that is why the very first English textbook of Mendelian genetics concluded with the thought that \u201cpermanent progress is a question of breeding rather than of pedagogics; a matter of gametes, not of training \u2026 the creature is not made but born\u201d (Punnett 1905, 60). <em>Translation: Not only do we now know a bit about how heredity works, but it\u2019s also the most important thing about you. Trust me, I\u2019m a scientist.<\/em><\/p>\n<p class=\"import-Normal\">Yet evolution is about how descendants come to differ from ancestors. Do we really know that your heredity, your DNA, your ancestry, is the most important thing about you? That you were born, not made? After all, we do know that you could be born into slavery or as a peasant, and come from a long line of enslaved people or peasants, and yet not have slavery or peasantry be the most important thing about you. Whatever your ancestors were may unfortunately constrain what you can become, but as a moral precept, it should not. But just as science is not purely \u201cfacts and objectivity,\u201d ancestry is not a strictly biological concept. Human ancestry is biopolitics, not biology.<\/p>\n<p class=\"import-Normal\">Evolution is fundamentally a theory about ancestry, and yet ancestors are, in the broad anthropological sense, sacred: ancestors are often more meaningful symbolically than biologically. Just a few years after <em>The Origin of Species <\/em>(Darwin 1859), the British politician and writer Benjamin Disraeli declared himself to be on the side of the angels, not the apes, and to \u201crepudiate with indignation and abhorrence those new-fangled theories\u201d (Monypenny, Flavelle, and Buckle 1920, 105). He turned his back on an ape ancestry and looked to the angel; yet, he did so as a prominent Jew-turned-Anglican, who had personally transcended his humble roots and risen to the pinnacle of the Empire. Ancestry was certainly important, and Disraeli was famously proud of his, but it was also certainly not the most important thing, not the primary determinant of his place in the world. Indeed, quite the opposite: Disraeli\u2019s life was built on the transcendence of many centuries of Jewish poverty and oppression in Europe. Humble ancestry was there to be superseded and nobility was there to be earned; Disraeli would later become the Earl of Beaconsfield. Clearly, \u201care you just your ancestry\u201d is not a value-neutral question, and \u201cthe creature is not made, but born\u201d is not a value-neutral answer.<\/p>\n<p class=\"import-Normal\">Ancestry being the most important thing about a person became a popular idea twice in 20th century science. First, at the beginning of the century, when the <strong>eugenics<\/strong> movement in America called attention to \u201cfeeble-minded stocks,\u201d which usually referred to the poor or to immigrants (see Figure 17.4; and see Chapter 2). This movement culminated in Congress restricting the immigration of \u201cfeeble-minded races\u201d (said to include Jews and Italians) in 1924, and the Supreme Court declaring it acceptable for states to sterilize their \u201cfeeble-minded\u201d citizens involuntarily in 1927. After the Nazis picked up and embellished these ideas during World War II, Americans moved swiftly away from them in some contexts (e.g., for most people of European descent) while still strictly adhering in other contexts (e.g., Japanese internment camps and immigration restrictions).<\/p>\n<figure style=\"width: 374px\" class=\"wp-caption alignright\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image4-6.png\" alt=\"Historic photo. People sit in front of a structure with a \u201cEugenic and Health Exhibit&quot; banner.\" width=\"374\" height=\"262\" \/><figcaption class=\"wp-caption-text\">Figure 17.4: Eugenic and Health Exhibit, Fitter Families exhibit, and examination building, Kansas State Free Fair. Credit: <a href=\"https:\/\/www.dnalc.org\/view\/16328-Gallery-14-Eugenics-Exhibit-at-the-Kansas-State-Free-Fair-1920.html\">Gallery 14: Eugenics Exhibit at the Kansas State Free Fair, 1920 ID (ID 16328)<\/a> by <a href=\"https:\/\/www.dnalc.org\/\">Cold Spring Harbor<\/a> (Courtesy American Philosophical Society) is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/3.0\/us\/\">CC BY-NC-ND 3.0 License.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\">Ancestry again became paramount in the drumming up of public support for the Human Genome Project in the 1990s. Public support for sequencing the human genome was encouraged by a popular science campaign that featured books titled <em>The Book of Man <\/em>(Bodmer and McKie 1997), <em>The Human Blueprint <\/em>(Shapiro 1991), and <em>The Code of Codes<\/em> (Kevles and Hood 1993). These books generally promised cures for genetic diseases and a deeper understanding of the human condition. We can certainly identify progress in molecular genetics over the last couple of decades since the human genome was sequenced, but that progress has notably not been accompanied by cures for genetic diseases, nor by deeper understandings of the human condition.<\/p>\n<p class=\"import-Normal\">Even at the most detailed and refined levels of genetic analysis, we still don\u2019t have much of an understanding of the actual basis by which things seem to \u201crun in families.\u201d While the genetic basis of simple, if tragic, genetic diseases have become well-known\u2014such as sickle-cell anemia, cystic fibrosis, and Tay-Sachs\u2019 Disease\u2014we still haven\u2019t found the ostensible genetic basis for traits that are thought to have a strong genetic component. For example, a recent genetic summary found over 12,000 genetic sites that contributed to height yet still explained only about 40-50 percent of the variation in height among European ancestry but no more than 10-20 percent of variation of other ancestries, which we know strongly runs in families (Yengo et al. 2022).<\/p>\n<p class=\"import-Normal\">Partly in reaction to the reductionistic hype of the Human Genome Project, the study of epigenetics has become the subject of great interest. One famous natural experiment involves a Nazi-imposed famine in Holland over the winter of 1944\u20131945. Children born during and shortly after the famine experienced a higher incidence of certain health problems as adults, many decades later. Apparently, certain genes had been down-regulated early in development and remained that way throughout the course of life. Indeed, this modified regulation of the genes in response to the severe environmental conditions may have been passed on to their children.<\/p>\n<p class=\"import-Normal\">Obviously one\u2019s particular genetic constitution may play an important role in one\u2019s life trajectory. But overvaluing that role may have important social and political consequences. In the first place, genotypes are rendered meaningful in a cultural universe. Thus, if you live in a strongly patriarchal society and are born without a Y chromosome (since human males are chromosomally XY and females XX), your genotype will indeed have a strong effect upon your life course. So even though the variation is natural, the consequences are political. The mediating factors are the cultural ideas about how people of different sexes ought to be treated, and the role of the state in permitting certain people to develop and thrive. More broadly, there are implications for public education if variation in intelligence is genetic. There are implications for the legal system if criminality is genetic. There are implications for the justice system if sexual preference, or sexual identity, is genetic. There are implications for the development of sports talent if that is genetic. And yet, even for the human traits that are more straightforward to measure and known to be strongly heritable, the DNA base sequence variation seems to explain little.<\/p>\n<p class=\"import-Normal\">Genetic determinism or <strong>hereditarianism<\/strong> is the idea that \u201cthe creature is made, not born\u201d\u2014or, in a more recent formulation by James Watson, that \u201cour fate is in our genes.\u201d One of the major implications drawn from genetic determinism is that the feature in question must inevitably express itself; therefore, we can\u2019t do anything about it. Therefore, we might as well not fund the social programs designed to ameliorate economic inequality and improve people\u2019s lives, because their courses are fated genetically. And therefore, they don\u2019t deserve better lives.<\/p>\n<p class=\"import-Normal\">All of the \u201ctherefores\u201d in the preceding paragraph are open to debate. What is important is that the argument relies on a very narrow understanding of the role of genetics in human life, and it misdirects the causes of inequality from cultural to natural processes. By contrast, instead of focusing on genes and imagining them to place an invisible limit upon social progress, we can study the ways in which your DNA sequence does <em>not<\/em> limit your capability for self-improvement or fix your place in a social hierarchy. In general, two such avenues exist. First, we can examine the ways in which the human body responds and reacts to environmental variation: human adaptability and plasticity. This line of research began with the anthropometric studies of immigrants by Franz Boas in the early 20th century and has now expanded to incorporate the epigenetic inheritance of modified human DNA. And second, we can consider how human lives are shaped by social histories\u2014especially the structural inequalities within the societies in which they grow up.<\/p>\n<p class=\"import-Normal\">Although it arises and is refuted every generation, the radical hereditarian position (genetic determinism) perennially claims to speak for both science and evolution. It does not. It is the voice of a radical fringe\u2014perhaps naive, perhaps evil. It is not the authentic voice of science or of evolution. Indeed, keeping Charles Darwin\u2019s name unsullied by protecting it from association with bad science often seems like a full-time job. Culture and epigenetics are very much a part of the human condition, and their roles are significant parts of the complete story of human evolution.<\/p>\n<p><span style=\"background-color: #00ffff\"><span style=\"text-decoration: underline\">(Sterilization of Indigenous women in Canada)<\/span> (https:\/\/www.thecanadianencyclopedia.ca\/en\/article\/sterilization-of-indigenous-women-in-canada)\u00a0<\/span><\/p>\n<h2 class=\"import-Normal\">Adaptationism and the Panglossian Paradigm<\/h2>\n<p class=\"import-Normal\">The story of human evolution, and the evolution of all life for that matter, is never settled because evolution is ongoing. Additionally, because the conditions that shape evolutionary trajectories are not predetermined, evolution itself is emergent. Even during periods of ecological stability, when fewer macroevolutionary changes occur, populations of organisms continue to experience change. When ecological stability is disrupted, populations must adapt to the changes. Darwin explained in naturalistic terms how animals adapt to their environments: traits that contribute to an organism's ability to survive and reproduce in specific environments will become more common. The most \u201cfit\u201d\u2014those organisms best suited to the <em>current<\/em> environmental conditions in which they live\u2014have survived over eons of the history of life on earth to cocreate ecosystems full of animals and plants. Our own bodies are full of evident adaptations: eyes for seeing, ears for hearing, feet for walking on, and so forth.<\/p>\n<p class=\"import-Normal\">But what about hands? Feet are adapted to be primarily weight-bearing structures (rather than grasping structures, as in the apes) and that is what we primarily use them for. But we use our hands in many ways: for fine-scale manipulation, greeting, pointing, stimulating a sexual partner, writing, throwing, and cooking, among other uses. So which of these uses express what hands are \u201cfor,\u201d when all of them express what hands do?<\/p>\n<p class=\"import-Normal\">Gould and Lewontin (1979) illustrate the problem with assuming that the function of a trait defines its evolutionary cause. Consider the case of Dr. Pangloss\u2014the protagonistic of Voltaire\u2019s <em>Candide<\/em>\u2014who believed that we lived in the best of all possible worlds. Gould and Lewontin use his pronouncement that \u201cnoses were made for spectacles and so we have spectacles\u201d to demonstrate the problem with assuming any trait has evolved for a specific purpose. Identifying a function of a trait does not necessitate that the function is the ultimate cause of the trait. Individual traits are not under selection pressures in isolation; in fact, an entire organism must be able to survive and reproduce in their environment. When natural selection results in adaptations, changes that occur in some traits can have cascading effects throughout the phenotype and features that are not under selection pressure can also change.<\/p>\n<figure style=\"width: 279px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image3-5.png\" alt=\"Human hand is smaller with smaller fingers and smoother skin compared to a chimpanzee hand.\" width=\"279\" height=\"264\" \/><figcaption class=\"wp-caption-text\">Figure 17.5: Drawings of a human hand (left) and a chimpanzee hand (right). Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__\/\">Human and chimpanzee hand (Figure 2.16)<\/a> by Mary Nelson original to <a href=\"https:\/\/explorations.americananthro.org\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">There is an important lesson in recognizing that what things do in the present is not a good guide to understanding why they came to exist. Gunpowder was invented for entertainment\u2014only later was it adopted for killing people. The Internet was invented to decentralize computers in case of a nuclear attack\u2014and only later adopted for social media. Apes have short thumbs and use their hands in locomotion; our ancestors stopped using their hands in locomotion by about six million years ago and had fairly modern-looking hands by about two million years ago. We can speculate that a combination of selection for abstract thought and dexterity led to evolution of the human hand, with its capability for toolmaking that exceeds what apes can do (see Figure 17.5). But let\u2019s face it\u2014how many tools have you made today?<\/p>\n<p class=\"import-Normal\">Consequently, we are obliged to see the human foot as having a purpose to which it is adapted and the human hand as having multiple purposes, most of which are different from what it originally evolved for. Paleontologists Gould and Elisabeth Vrba suggested that an original use be regarded as an adaptation and any additional uses be called \u201c<strong>exaptations.<\/strong>\u201d Thus, we would consider the human hand to be an adaptation for toolmaking and an exaptation for writing. So how do we know whether any particular feature is an adaptation, like the walking foot, rather than an exaptation, like the writing hand? Or more broadly, how can we reason rigorously from what a feature does to what it evolved for?<\/p>\n<p class=\"import-Normal\">The answer to the question \u201cwhat did this feature evolve for?\u201d creates an origin myth. This origin myth contains three assumptions: (1) features can be isolated as evolutionary units; (2) there is a specific reason for the existence of any particular feature; and (3) there is a clear and simplistic explanation for why the feature evolved.<\/p>\n<figure style=\"width: 378px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image9-8.png\" alt=\"Head with images and human qualities drawn on it. Journal title printed at the bottom.\" width=\"378\" height=\"437\" \/><figcaption class=\"wp-caption-text\">Figure 17.6: According to the early 19th century science of phrenology, units of personality could be mapped onto units in the head, as shown on this cover of The Phrenology Journal. Credit: <a href=\"https:\/\/wellcomecollection.org\/works\/b6skynug\">Phrenology; Chart<\/a> [slide number 5278, photo number: L0000992, original print from Dr. E. Clark, The Phrenological Journal (Know Thyself)] by <a href=\"https:\/\/wellcomecollection.org\/\">Wellcome Collection<\/a>, is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/legalcode\">CC BY 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">The first assumption was appreciated a century ago as the \u201cunit-character problem.\u201d Are the units by which the body grows and evolves the same as units we name? This is clearly not the case: we have genes and we have noses, and we have genes that affect noses, but we don\u2019t have \u201cnose genes.\u201d What is the relationship between the evolving elements that we see, identify, and name, and the elements that biologically exist and evolve? It is hard to know, but we can use the history of science as a guide to see how that fallacy has been used by earlier generations. Back in the 19th century, the early anatomists argued that since the brain contained the mind, they could map different mental states (acquisitiveness, punctuality, sensitivity) onto parts of the brain. Someone who was very introspective, say, would have an enlarged introspection part of the brain, a cranial bulge to represent the hyperactivity of this mental state. The anatomical science was known as <strong>phrenology<\/strong>, and it was predicated on the false assumption that units of thought or personality or behavior could be mapped to distinct parts of the brain and physically observed (see Figure17.6). This is the fallacy of reification, imagining that something named is something real.<\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Long alt text: Side view of human head. At the top are the words \u201cKnow Thyself.\u201d On the upper head are small illustrations and word qualities such as \u201cfriendship,\u201d \u201cself-esteem,\u201d and \u201csecretiveness.\u201d On the lower part of the man\u2019s man\u2019s face are the words <em>The Phrenological Journal and Science of Health, A First Class Monthly<\/em>. The caption at the bottom reads: \u201cSpecially devoted to the \u2018.\u2019 Contains PHRENOLOGY and PHYSIOGNOMY, with all the SIGNS OF CHARACTER, and how to read them; ETHNOLOGY, or the Natural History of Man in all his relations.\u201d (All emphases in original.)<\/span><\/p>\n<figure style=\"width: 295px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image1-8-1.png\" alt=\"A black-and-white drawing of a chimpanzee head and face.\" width=\"295\" height=\"236\" \/><figcaption class=\"wp-caption-text\">Figure 17.7: Chimpanzees have big ears. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Chimpanzee_head_sketch.png\">Chimpanzee head sketch<\/a> by <a href=\"https:\/\/de.wikipedia.org\/wiki\/Benutzer:Roger_Zenner\">Roger Zenner<\/a>, original by Brehms Tierleben (1887), is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">The second assumption, that everything has a reason, has long been recognized as a core belief of religion. Our desire to impose order and simplicity on the workings of the universe, however, does not constrain it to obey simple and orderly causes. Magic, witchcraft, spirits, and divine agency are all powerful explanations for why things happen. Consequently, it is probably not a good idea to lump natural selection in with those. Sometimes things do happen for a reason, of course, but other times things happen as byproducts of other things, or for very complicated and entangled reasons, or for no reason at all. What phenomena have reasons and thereby merit explanation? Chimpanzees have very large testicles, and we think we know why: their promiscuous sexual behavior triggers intense competition for high sperm count. But chimpanzees also have very large ears, but much less scientific attention has been paid to this trait (see Figure 17.7). Why not? Why should there be a reason for chimp testicles but not for chimp ears? What determines the kinds of features that we try to explain, as opposed to the ones that we do not? Again, the assumption that any specific feature has a reason is metaphysical; that is to say, it may be true in any particular case, but to assume it in all cases is gratuitous.<\/p>\n<p class=\"import-Normal\">And third, the possibility of knowing what the reason for any particular feature is, assuming that it has one, is a challenge for evolutionary epistemology (the theory of how we know things). Consider the big adaptations of our lineage: bipedalism and language. Nobody doubts that they are good, and they evolved by natural selection, and we know how they work. But why did they evolve? If talking and walking are simply better than not talking and not walking, then why did they evolve in just a single branch of the ape lineage in the primate family tree? We don\u2019t know what bipedalism evolved for, although there are plenty of speculations: walking long distances, running long distances, cooling the head, seeing over tall grass, carrying babies, carrying food, wading, threatening, counting calories, sexual display, and so on. Neither do we know what language evolved for, although there are speculations: coordinating hunting, gossiping, manipulating others. But it is also possible that bipedality is simply the way that a small arboreal ape travels on the ground, if it isn\u2019t in the treetops. Or that language is simply the way that a primate with small canine teeth and certain mental propensities comes to communicate. If that were true, then there might be no reason for bipedality or language: having the unique suite of preconditions and a fortuitous set of circumstances simply set them in motion, and natural selection elaborated and explored their potentials. It is possible that walking and talking simply solved problems that no other lineage had ever solved; but even if so, the fact remains that the rest of the species in the history of life have done pretty well without having solved them.<\/p>\n<p class=\"import-Normal\">It is certainly very optimistic to think that all three assumptions (that organisms can be meaningfully atomized, that everything has a reason, and that we can know the reason) would be simultaneously in effect. Indeed, just as there are many ways of adapting (genetically, epigenetically, behaviorally, culturally), there are also many ways of being nonadaptive, which would imply that there is no reason at all for the feature in question.<\/p>\n<p class=\"import-Normal\">First, there is the element of randomness of population histories. There are more cases of sickle-cell anemia among sub-Saharan Africans than other peoples, and there is a reason for it: carriers of sickle-cell anemia have a resistance to malaria, which is more frequent in parts of Africa (as discussed in Chapters 4 and 14). But there are more cases of a blood disease called variegated porphyria, a rare genetic metabolic disorder, in the Afrikaners of South Africa (descendants of mostly Dutch settlers in the 17th century) than in other peoples, and there is no reason for it. Yet we know the cause: One of the founding Dutch colonial settlers had the <strong>allele<\/strong>\u2013a variant of a gene\u2013and everyone in South Africa with it today is her descendant. But that is not a reason\u2014that is simply an accident of history.<\/p>\n<p class=\"import-Normal\">Second, there is the potential mismatch between the past and the present. The value of a particular feature in the past may be changed as the environmental circumstances change. Our species is diurnal, and our ancestors were diurnal. But beginning around a few hundred thousand years ago, our ancestors could build fires, which extended the light period, which was subsequently further amplified by lamps and candles. And over the course of the 20th century, electrical power has made it possible for people to stay up very late when it is dark\u2014working, partying, worrying\u2014to a greater extent than any other closely related species. In other words, we evolved to be diurnal, yet we are now far more nocturnal than any of our recent ancestors or close relatives. Are we adapting to nocturnality? If so, why? Does it even make any sense to speak of the human occupation of a nocturnal ape niche, despite the fact that we empirically seem to be doing just that? And if so, does it make sense to ask what the reason for it is?<\/p>\n<p class=\"import-Normal\">Third, there is a genetic phenomenon known as a selective sweep, or the hitchhiker effect. Imagine three genes\u2014A, B, and C\u2014located very closely together on a chromosome. They each have several variants, or alleles, in the population. Now, for whatever reason, it becomes beneficial to have one of the B alleles, say B4; this B4 allele is now under strong positive selection. Obviously, we will expect future generations to be characterized by mostly B4. But what was B4 attached to? Because whatever A and C alleles were adjacent to it will also be quickly spread, simply by virtue of the selection for B4. Even if the A and C alleles are not very good, they will spread because of the good B4 allele between them. Eventually the linkage groups will break up because of genetic crossing-over in future generations. But in the meantime, some random version of genes A and C are proliferating in the species simply because they are joined to superior allele B4. And clearly, the A and C alleles are there because of selection\u2014but not because of selection <em>for<\/em> them!<\/p>\n<p class=\"import-Normal\">Fourth, some features are simply consequences of other properties rather than adaptations to external conditions. We already noted the phenomenon of allometric growth, in which some physical features have to outgrow others to maintain function at an increased size. Can we ask the reason for the massive brow ridges of <em>Homo erectus<\/em>, or are brow ridges simply what you get when you have a conjunction of thick skull bones, a large face, and a sloping forehead\u2014and, thus, again would have a cause but no reason?<\/p>\n<p class=\"import-Normal\">Fifth, some features may be underutilized and on the way out. What is the reason for our two outer toes? They aren\u2019t propulsive, they don\u2019t do anything, and sometimes they\u2019re just in the way. Obviously they are there because we are descended from ancestors with five digits on their hands and feet. Is it possible that a million years from now, we will just have our three largest toes, just as the ancestors of the horse lost their digits in favor of a single hoof per limb? Or will our outer toes find another use, such as stabilizing the landings in our personal jet-packs? For the time being, we can just recognize vestigiality as another nonadaptive explanation for the presence of a given feature.<\/p>\n<p class=\"import-Normal\">Finally, Darwin himself recognized that many obvious features do not help an animal survive. Some things may instead help an animal breed. The peacock\u2019s tail feathers do not help it eat, but they do help it mate. There is competition, but only against half of the species. Darwin called this <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1084\">sexual selection<\/a><\/strong>. Its result is not a fit to the environment but, rather, a fit to the opposite sex. In some species, that is literally the case, as the male and female genitalia have specific ways of anatomically fitting together. The specific form is less important than the specific match, so inquiring about the reason for a particular form of the reproductive anatomy may be misleading. The specific form may be effectively random, as long as it fits the opposite sex and is different from the anatomies of other species. Nor is sexual selection the only form of selection that can affect the body differently from natural selection. Competition might also take place between biological units other than organisms\u2014perhaps genes, perhaps cells, or populations, or species. The spread of cultural things, such as head-binding or cheap refined fructose or forced labor, can have significant effects upon bodies, which are also not adaptations produced by natural selection. They are often adaptive physiological responses to stresses but not the products of natural selection.<\/p>\n<p class=\"import-Normal\">With so many paths available by which a physical feature might have organically arisen without having been the object of natural selection, it is unwise to assume that any individual trait is an adaptation. And that generalization applies to the best-known, best-studied, and most materially based evolutionary adaptations of our lineage. But our cultural behaviors are also highly adaptive, so what about our most familiar social behaviors? Patriarchy, hierarchy, warfare\u2014are these adaptations? Do they have reasons? Are they good for something?<\/p>\n<p class=\"import-Normal\">This is where some sloppy thinking has been troublesome. What would it mean to say that patriarchy evolved by natural selection in the human species? If, on the one hand, it means that the human mind evolved by natural selection to be able to create and survive in many different kinds of social and political regimes, of which patriarchy is one, then biological anthropologists will readily agree. If, on the other hand, it means that patriarchy evolved by natural selection, that implies that patriarchy is genetically determined (since natural selection is a genetic process) and out-reproduced the alleles for other, more egalitarian, social forms. This in turn would imply that patriarchy is an adaptation and therefore of some beneficial value in the past and has become an ingrained part of human nature today. This would be bad news, say, if you harbored ambitions of dismantling it. Dismantling patriarchy in that case would be to go against nature, a futile gesture. In other words, this latter interpretation would be a naturalistic manifesto for a conservative political platform: don\u2019t try to dismantle the patriarchy, because it is within us, the product of evolution\u2014suck it up and live with it.<\/p>\n<p class=\"import-Normal\">Here, evolution is being used as a political instrument for transforming the human genome into an imaginary glass ceiling against equality. There is thus a convergence between the pseudo-biology of crude <strong>adaptationism <\/strong>(the idea that everything is the product of natural selection) and the pseudo-biology of hereditarianism. Naturalizing inequality is not the business of evolutionary theory, and it represents a difficult moral position for a scientist to adopt, as well as a poor scientific position.<\/p>\n<div class=\"textbox shaded\">\n<p>Dig Deeper: Evolution of Humans and Its Effects (to be reviewed)<\/p>\n<p>As humans have caused the emergence of the Anthropocene, it is important to inform scholars about the effect of our social and cultural evolution on the rest of the world. Richard Robbins\u2019 <em>Global Problems and Culture of Capitalism<\/em> explains how the modern culture of consumption has been extremely successful at accommodating populations of people far larger than previously possible. Robbins claims that the globalization attributed to capitalism has allowed the world to make full use of its environmental resources, providing necessities and innovative technologies to humans all over the world (Robbins &amp; Dowty, 2019). In other words, capitalism is an anthropocentric cultural system that highly benefits humans and facilitates our survival with little regard to the development and survival of other forms of life. It would be of high relevance to introduce the idea that our cultural evolution and capacity to modify the environment to meet our needs have established new environmental conditions in which the human species' survival and reproduction rate expand at the detriment of ecosystems and endangerment of other primates and non-human species.<\/p>\n<p>According to a 2019 UN report, following the 16th century, the world has entered a period of extreme environmental destruction that is generating ecological modifications and has led to the extinction of at least 680 vertebrate species and over 9 percent of the domesticated mammals used for food and agriculture (United Nations, 2019). Human lifestyles are causing changes that\u2014if not taken into consideration\u2014could lead to our extinction as a species. The recognition that our evolutionary behavioural development is causing environmental destruction may be the first step for our species to take accountability for the damage that it is causing to others and prevent further damage.<\/p>\n<\/div>\n<p>&nbsp;<\/p>\n<h2 class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Concluding Thoughts<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Now that you have finished reading this chapter, you are equipped to understand the historical and political dimensions of evolution. Evolution is an ongoing process of change and diversification. Evolutionary theory is a tool that we use to understand this process. The development of evolutionary theory is shaped both by scientific innovation and political engagement. Since Darwin first articulated natural selection as an observable mechanism by which species adapt to their environments, our understanding of evolution has grown. Initially, scientists focused on the adaptive aspects of evolution. However, with the emergence of genetics, our understanding of heredity and the level at which evolution acts has changed. Genetics led to a focus on the molecular dimensions of evolution. For some, this focus resulted in reductive accounts of evolution. Further developments in our understanding of evolution shifted our view to epigenetic processes and how organisms shape their own evolutionary pressures (e.g., niche construction).<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Evolutionary theory will continue to develop in the future as we invent new technologies, describe new dimensions of biology, and experience cultural changes. Current innovations in evolutionary theory are asking us to consider evolutionary forces beyond natural selection and genetics to include the ways organisms shape their environments (niche construction), inheritances beyond genetics (inclusive inheritance), constraints on evolutionary change (developmental bias), and the ability of bodies to change in response to external factors (plasticity). The future of evolutionary theory looks bright as we continue to explore these and other dimensions. Biological anthropology is well-positioned to be a lively part of this conversation, as it extends standard evolutionary theory by considering the role of culture, social learning, and human intentionality in shaping the evolutionary trajectories of humans (Zeder 2018). Remember, at root, human evolutionary theory consists of two propositions: (1) the human species is descended from other similar species and (2) natural selection has been the primary agent of biological adaptation. Pretty much everything else is subject to some degree of contestation.<\/span><\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Review Questions<\/h2>\n<ul>\n<li class=\"import-Normal\">How is the study of your ancestors biopolitical, not just biological? Does that make it less scientific or differently scientific?<\/li>\n<li class=\"import-Normal\">What was gained by reducing organisms to genotypes and species to gene pools? What is gained by reintroducing bodies and species into evolutionary studies?<\/li>\n<li class=\"import-Normal\">How do genetic or molecular studies complement anatomical studies of evolution?<\/li>\n<li class=\"import-Normal\">How are you reducible to your ancestry? If you could meet your ancestors from the year 1700 (and you would have well over a thousand of them!), would their lives be meaningfully similar to yours? Would you even be able to communicate with them?<\/li>\n<li class=\"import-Normal\">The molecular biologist Fran\u00e7ois Jacob argued that evolution is more like a tinkerer than an engineer. In what ways do we seem like precisely engineered machinery, and in what ways do we seem like jerry-rigged or improvised contraptions?<\/li>\n<li class=\"import-Normal\">How might biological anthropology contribute to future developments in evolutionary theory?<\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\">Key Terms<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\"><strong>Adaptation<\/strong>: A fit between the organism and environment.<\/p>\n<p class=\"import-Normal\"><strong>Adaptationism<\/strong>: The idea that everything is the product of natural selection.<\/p>\n<p class=\"import-Normal\"><strong>Allele<\/strong>: A genetic variant.<\/p>\n<p class=\"import-Normal\"><strong>Allometry<\/strong>: The differential growth of body parts.<\/p>\n<p class=\"import-Normal\"><strong>Canalization<\/strong>: The tendency of a growing organism to be buffered toward normal development.<\/p>\n<p class=\"import-Normal\"><strong>Epigenetics<\/strong>: The study of how genetically identical cells and organisms (with the same DNA base sequence) can nevertheless differ in stably inherited ways.<\/p>\n<p class=\"import-Normal\"><strong>Eugenics<\/strong>: An idea that was popular in the 1920s that society should be improved by breeding \u201cbetter\u201d kinds of people.<\/p>\n<p class=\"import-Normal\"><strong>Evo-devo<\/strong>: The study of the origin of form; a contraction of \u201cevolutionary developmental biology.\u201d<\/p>\n<p class=\"import-Normal\"><strong>Exaptation<\/strong>: An additional beneficial use for a biological feature.<\/p>\n<p class=\"import-Normal\"><strong>Extinction<\/strong>: The loss of a species from the face of the earth.<\/p>\n<p class=\"import-Normal\"><strong>Gene<\/strong>: A stretch of DNA with an identifiable function (sometimes broadened to include any DNA with recognizable structural features as well).<\/p>\n<p class=\"import-Normal\"><strong>Gene pool<\/strong>: Hypothetical summation of the entire genetic composition of population or species.<\/p>\n<p class=\"import-Normal\"><strong>Genotype<\/strong>: Genetic constitution of an individual organism.<\/p>\n<p class=\"import-Normal\"><strong>Hereditarianism<\/strong>: The idea that genes or ancestry is the most crucial or salient element in a human life. Generally associated with an argument for natural inequality on pseudo-genetic grounds.<\/p>\n<p class=\"import-Normal\"><strong>Hox genes<\/strong>: A group of related genes that control for the body plan of an embryo along the head-tail axis.<\/p>\n<p class=\"import-Normal\"><strong>Inheritance of acquired characteristics<\/strong>: The idea that you pass on the features that developed during your lifetime, not just your genes; also known as Lamarckian inheritance.<\/p>\n<p class=\"import-Normal\"><strong>Natural selection<\/strong>: A consistent bias in survival and fertility, leading to the overrepresentation of certain features in future generations and an improved fit between an average member of the population and the environment.<\/p>\n<p class=\"import-Normal\"><strong>Niche construction<\/strong>: The active engagement by which species transform their surroundings in favorable ways, rather than just passively inhabiting them.<\/p>\n<p class=\"import-Normal\"><strong>Phenotype<\/strong>: Observable manifestation of a genetic constitution, expressed in a particular set of circumstances. The suite of traits of an organism.<\/p>\n<p class=\"import-Normal\"><strong>Phrenology<\/strong>: The 19th-century anatomical study of bumps on the head as an indication of personality and mental abilities.<\/p>\n<p class=\"import-Normal\"><strong>Plasticity<\/strong>: The tendency of a growing organism to react developmentally to its particular conditions of life.<\/p>\n<p class=\"import-Normal\"><strong>Punctuated equilibria<\/strong>: The idea that species are stable through time and are formed very rapidly relative to their duration. (The opposite theory, that species are unstable and constantly changing through time, is called phyletic gradualism.)<\/p>\n<p class=\"import-Normal\"><strong>Scientific racism<\/strong>: The use of pseudoscientific evidence to support or legitimize racial hierarchy and inequality.<\/p>\n<p class=\"import-Normal\"><strong>Sexual selection<\/strong>: Natural selection arising through preference by one sex for certain characteristics in individuals of the other sex.<\/p>\n<p class=\"import-Normal\"><strong>Species selection<\/strong>: A postulated evolutionary process in which selection acts on an entire species population, rather than individuals.<\/p>\n<h2 class=\"import-Normal\">About the Authors<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image2-4-1.jpg\" alt=\"A bearded man wearing glasses smiles at the camera. \" width=\"202\" height=\"218\" \/><\/p>\n<h3 class=\"import-Normal\"><strong>Jonathan Marks, Ph.D.<\/strong><\/h3>\n<p class=\"import-Normal\">University of North Carolina at Charlotte, <a class=\"rId41\" href=\"mailto:jmarks@uncc.edu\">jmarks@uncc.edu<\/a><\/p>\n<p class=\"import-Normal\">Jonathan Marks is Professor of Anthropology at the University of North Carolina at Charlotte. He has published many books and articles on broad aspects of biological anthropology. In 2006 he was elected a Fellow of the American Association for the Advancement of Science. In 2012 he was awarded the First Citizen\u2019s Bank Scholar\u2019s Medal from UNC Charlotte. In recent years he has been a Visiting Research Fellow at the ESRC Genomics Forum in Edinburgh, a Visiting Research Fellow at the Max Planck Institute for the History of Science in Berlin, and a Templeton Fellow at the Institute for Advanced Study at Notre Dame. His work has received the W. W. Howells Book Prize and the General Anthropology Division Prize for Exemplary Cross-Field Scholarship from the American Anthropological Association as well as the J. I. Staley Prize from the School for Advanced Research. Two of his books are titled <em>What It Means to Be 98% Chimpanzee<\/em> and <em>Why I Am Not a Scientist<\/em>, but actually he is about 98 percent scientist and not a chimpanzee.<\/p>\n<p class=\"import-Normal\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image10-3.jpg\" alt=\"A bearded man wearing a fedora hat looks off in the distance. \" width=\"232\" height=\"232\" \/><\/p>\n<h3 class=\"import-Normal\"><strong>Adam P. Johnson, M.A.<\/strong><\/h3>\n<p class=\"import-Normal\">University of North Carolina at Charlotte\/University of Texas at San Antonio, <a class=\"rId43\" href=\"mailto:ajohn344@uncc.edu\">ajohn344@uncc.edu<\/a><\/p>\n<p class=\"import-Normal\">Adam Johnson is a doctoral candidate at the University of Texas at San Antonio and part-time lecturer at the University of North Carolina at Charlotte. He earned his M.A. in anthropology at UNC-Charlotte in 2017 and will complete his Ph.D. in anthropology at UTSA by 2024. His interests include human-animal relations, science studies, primate behavior, ecology, and the history of anthropology. His recent research project analyzes the social, historical, political, and evolutionary dimensions that shape human-javelina encounters. His goal is to understand how humans and animals find ways to get along in a precarious world.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration <strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\">Ackermann, Rebecca Rogers, Alex Mackay, and Michael L. Arnold. 2016. \u201cThe Hybrid Origin of \u2018Modern\u2019 Humans.\u201d <em>Evolutionary Biology<\/em> 43 (1): 1\u201311.<\/p>\n<p class=\"import-Normal\">Bateson, Patrick, and Peter Gluckman. 2011. <em>Plasticity, Robustness, Development and Evolution<\/em>. New York: Cambridge University Press.<\/p>\n<p class=\"import-Normal\">Cosans, Christopher E. 2009. <em>Owen's Ape and Darwin's Bulldog: Beyond Darwinism and Creationism<\/em>. Bloomington, IN: Indiana University Press.<\/p>\n<p class=\"import-Normal\">Desmond, Adrian, and James Moore. 2009. <em>Darwin's Sacred Cause: How a Hatred of Slavery Shaped Darwin's Views on Human Evolution<\/em>. New York: Houghton Mifflin Harcourt.<\/p>\n<p class=\"import-Normal\">Dobzhansky, Theodosius, Francisco J. Ayala, G. Ledyard Stebbins, and James W. Valentine. 1977. <em>Evolution<\/em>. San Francisco: W.H. Freeman and Company.<\/p>\n<p class=\"import-Normal\">Fuentes, Agust\u00edn. 2017. <em>The Creative Spark: How Imagination Made Humans Exceptional<\/em>. New York: Dutton.<\/p>\n<p class=\"import-Normal\">Gould, Stephen J. 2003.<em> The Structure of Evolutionary Theory<\/em>. Cambridge, MA: Harvard University Press.<\/p>\n<p class=\"import-Normal\">Haraway, Donna J. 1989. <em>Primate Visions: Gender, Race, and Nature in the World of Modern Science<\/em>. New York: Routledge.<\/p>\n<p class=\"import-Normal\">Huxley, Thomas. 1863. <em>Evidence as to Man's Place in Nature<\/em>. London: Williams &amp; Norgate.<\/p>\n<p class=\"import-Normal\">Jablonka, Eva, and Marion J. Lamb. 2005. <em>Evolution in Four Dimensions: Genetic, Epigenetic, Behavioral, and Symbolic Variation in the History of Life<\/em>. Cambridge, MA: The MIT Press.<\/p>\n<p class=\"import-Normal\">Kuklick, Henrika, ed. 2008. <em>A New History of Anthropology<\/em>. New York: Blackwell.<\/p>\n<p class=\"import-Normal\">Laland, Kevin N., Tobias Uller, Marcus W. Feldman, Kim Sterelny, Gerd B. Muller, Armin Moczek, Eva Jablonka, and John Odling-Smee. 2015. \u201cThe Extended Evolutionary Synthesis: Its Structure, Assumptions and Predictions.\u201d <em>Proceedings of the Royal Society, Series B<\/em> 282 (1813): 20151019.<\/p>\n<p class=\"import-Normal\">Lamarck, Jean Baptiste. 1809. <em>Philosophie Zoologique<\/em>. Paris: Dentu.<\/p>\n<p class=\"import-Normal\">Landau, Misia. 1991. <em>Narratives of Human Evolution<\/em>. New Haven: Yale University Press.<\/p>\n<p class=\"import-Normal\">Lee, Sang-Hee. 2017. <em>Close Encounters with Humankind: A Paleoanthropologist Investigates Our Evolving Species<\/em>. New York: W. W. Norton.<\/p>\n<p class=\"import-Normal\">Livingstone, David N. 2008. <em>Adam's Ancestors: Race, Religion, and the Politics of Human Origins<\/em>. Baltimore: Johns Hopkins University Press.<\/p>\n<p class=\"import-Normal\">Marks, Jonathan. 2015. <em>Tales of the Ex-Apes: How We Think about Human Evolution<\/em>. Berkeley, CA: University of California Press.<\/p>\n<p class=\"import-Normal\">Pigliucci, Massimo. 2009. \u201cThe Year in Evolutionary Biology 2009: An Extended Synthesis for Evolutionary Biology.\u201d <em>Annals of the New York Academy of Sciences<\/em> 1168: 218\u2013228.<\/p>\n<p class=\"import-Normal\">Simpson, George Gaylord. 1949. <em>The Meaning of Evolution: A Study of the History of Life and of Its Significance for Man<\/em>. New Haven: Yale University Press.<\/p>\n<p class=\"import-Normal\">Sommer, Marianne. 2016.<em> History Within: The Science, Culture, and Politics of Bones, Organisms, and Molecules<\/em>. Chicago: University of Chicago Press.<\/p>\n<p class=\"import-Normal\">Stoczkowski, Wiktor. 2002. <em>Explaining Human Origins: Myth, Imagination and Conjecture<\/em>. New York: Cambridge University Press.<\/p>\n<p class=\"import-Normal\">Tattersall, Ian, and Rob DeSalle. 2019. <em>The Accidental Homo sapiens: Genetics, Behavior, and Free Will<\/em>. New York: Pegasus.<\/p>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\">Barton, Robert A. 1996. \"Neocortex Size and Behavioural Ecology in Primates.\" <em>Proceedings of the Royal Society of London. Series B: Biological Sciences<\/em> 263 (1367): 173\u2013177.<\/p>\n<p class=\"import-Normal\">Bodmer, Walter, and Robin McKie. 1997. <em>The Book of Man: The Hman Genome Project and the Quest to Discover our Genetic Heritage.<\/em> Oxford University Press.<\/p>\n<p class=\"import-Normal\">Darwin, Charles. 1859.<em> On the Origin of Species by Means of Natural Selection, or, the Preservation of Favoured Races in the Struggle for Life<\/em>. London: J. Murray.<\/p>\n<p class=\"import-Normal\">Darwin, Charles. 1871. <em>The Descent of Man, and Selection in Relation to Sex.<\/em> London: J. Murray.<\/p>\n<p class=\"import-Normal\">Dawkins, Richard. 1976. <em>The Selfish Gene. <\/em>Oxford University Press.<\/p>\n<p class=\"import-Normal\">Deacon, T. W. 1998. <em>The Symbolic Species: The Co-evolution of Language and the Brain<\/em>. W. W. Norton &amp; Company.<\/p>\n<p class=\"import-Normal\">Eldredge, N., and S. J. Gould. 1972. \"Punctuated Equilibria: An Alternative to Phyletic Gradualism.\" In <em>Models in Paleobiology<\/em>, edited by T. J. Schopf, 82\u2013115. San Francisco: W. H. Freeman.<\/p>\n<p class=\"import-Normal\">Gould, Stephen J. 2003.<em> The Structure of Evolutionary Theory<\/em>. Cambridge, MA: Harvard University Press.<\/p>\n<p class=\"import-Normal\">Gould, Stephen J. 1996. <em>Mismeasure of Man<\/em>. New York: WW Norton &amp; Company.<\/p>\n<p class=\"import-Normal\">Gould, Stephen Jay, and Richard C. Lewontin. 1979. \"The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme.\" <em>Proceedings of the Royal Society of London. Series B: Biological Sciences<\/em> 205 (1151): 581\u2013598.<\/p>\n<p class=\"import-Normal\">Haeckel, Ernst. 1868. <em>Nat\u00fcrliche Sch\u00f6pfungsgeschichte<\/em>. Berlin: Reimer.<\/p>\n<p class=\"import-Normal\">Huxley, Thomas Henry. 1863. <em>Evidence as to Man\u2019s Place in Nature. <\/em>London: Williams and Norgate.<\/p>\n<p class=\"import-Normal\">Kaufman, Thomas C., Mark A. Seeger, and Gary Olsen. 1990. \"Molecular and Genetic Organization of the Antennapedia Gene Complex of <em>Drosophila melanogaster<\/em>.\" <em>Advances in Genetics<\/em> 27: 309\u2013362.<\/p>\n<p class=\"import-Normal\">Kellogg, Vernon. 1917. <em>Headquarters Nights<\/em>. Boston: The Atlantic Monthly Press.<\/p>\n<p class=\"import-Normal\">Kevles, Daniel J., and Leroy Hood. 1993. <em>The Code of Codes: Scientific and Social Issues in the Human Genome Project<\/em>. Cambridge, MA: Harvard University Press.<\/p>\n<p class=\"import-Normal\">Lewontin, Richard, Steven Rose, and Leon Kamin. 2017. <em>Not in Our Genes\u202f: Biology, Ideology, and Human Nature<\/em>, 2nd ed. Chicago: Haymarket Books.<\/p>\n<p class=\"import-Normal\">Lloyd, Elisabeth A., and Stephen J. Gould. 1993. \"Species Selection on Variability.\" <em>Proceedings of the National Academy of Sciences<\/em> 90 (2): 595\u2013599.<\/p>\n<p class=\"import-Normal\">Marks, Jonathan. 2015. \u201cThe Biological Myth of Human Evolution.\u201d In <em>Biologising the Social Sciences: Challenging Darwinian and Neuroscience Explanations<\/em>, edited by David Canter and David A. Turner, 59\u201378. London: Routledge.<\/p>\n<p class=\"import-Normal\">Monypenny, William Flavelle, and George Earle Buckle. 1929. <em>The Life of Benjamin Disraeli, Earl of Beaconsfield, Volume II: 1860\u20131881<\/em>. London: John Murray.<\/p>\n<p class=\"import-Normal\">Potts, Rick. 1998. \u201cVariability Selection in Hominid Evolution.\u201d <em>Evolutionary Anthropology <\/em><em>7<\/em><em>:<\/em> 81\u201396.<\/p>\n<p class=\"import-Normal\">Punnett, R. C. 1905. <em>Mendelism<\/em>. Cambridge: Macmillan and Bowes.<\/p>\n<p class=\"import-Normal\">Shapiro, Robert. 1991. <em>The Human Blueprint: The Race to Unlock the Secrets of Our Genetic Script.<\/em> New York: St. Martin\u2019s Press.<\/p>\n<p class=\"import-Normal\">Shultz, Susanne, Emma Nelson, and Robin Dunbar. 2012. \"Hominin Cognitive Evolution: Identifying Patterns and Processes in the Fossil and Archaeological Record.\" <em>Philosophical Transactions of the Royal Society B: Biological Sciences<\/em> 367 (1599): 2130\u20132140.<\/p>\n<p class=\"import-Normal\">Spencer, Herbert. 1864. <em>Principles of Biology.<\/em> London: Williams and Norgate.<\/p>\n<p class=\"import-Normal\">Watson, James D. 1990. \"The Human Genome Project: Past, Present, and Future.\" <em>Science<\/em> 248 (4951): 44\u201349.<\/p>\n<p class=\"import-Normal\">Yengo, L., Vedantam, S., Marouli, E., Sidorenko, J., Bartell, E., Sakaue, S., Graff, M., Eliasen, A.U., Jiang, Y., Raghavan, S. and Miao, J., 2022. A saturated map of common genetic variants associated with human height. <em>Nature<\/em>, <em>610 <\/em>(7933): 704-712.<\/p>\n<p class=\"import-Normal\">Zeder, Melinda A. 2018. \"Why Evolutionary Biology Needs Anthropology: Evaluating Core Assumptions of the Extended Evolutionary Synthesis.\" <em>Evolutionary Anthropology: Issues, News, and Reviews<\/em> 27 (6): 267\u2013284.<\/p>\n<\/div>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1759\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1759\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\">Jonathan Marks, Ph.D., University of North Carolina at Charlotte<\/p>\n<p class=\"import-Normal\">Adam P. Johnson, M.A., University of North Carolina at Charlotte\/University of Texas at San Antonio<\/p>\n<p class=\"import-Normal\"><em>This chapter is an adaptation of \"<\/em><a class=\"rId9\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__\/\"><em>Chapter 2: Evolution<\/em><\/a><em>\u201d by Jonathan Marks. In <\/em><a class=\"rId10\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, first edition<\/em><\/a><em>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under <\/em><a class=\"rId11\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\"><em>CC BY-NC 4.0<\/em><\/a><em>. <\/em><\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\"><span style=\"color: #ffffff\">Learning Objectives<\/span><\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li>Explain the relationship among genes, bodies, and organismal change.<\/li>\n<li>Discuss the shortcomings of simplistic understandings of genetics.<\/li>\n<li>Describe what is meant by the \"biopolitics of heredity.\"<\/li>\n<li>Discuss issues caused by misuse of ideas about adaptations and natural selection.<\/li>\n<li>Examine and correct myths about evolution.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p class=\"import-Normal\">The Human Genome Project, an international initiative launched in 1990, sought to identify the entire genetic makeup of our species. For many scientists, it meant trying to understand the genetic underpinnings of what made humans uniquely human. James Watson, a codiscoverer of the helical shape of DNA, wrote that \u201cwhen finally interpreted, the genetic messages encoded within our DNA molecules will provide the ultimate answers to the chemical underpinnings of human existence\u201d (Watson 1990, 248). The underlying message is that what makes humans unique can be found in our <strong>genes<\/strong>. The Human Genome Project hoped to find the core of who we are and where we come from.<\/p>\n<p class=\"import-Normal\">Despite its lofty goal, the Human Genome Project\u2014even after publishing the entire human genome in January 2022\u2014could not fully account for the many factors that contribute to what it is to be human. Richard Lewontin, Steven Rose, and Leon Kamin (2017) argue that genetic determinism of the sort assumed by the Human Genome Project neglects other essential dimensions that contribute to the development and evolution of human bodies, not to mention the role that culture plays. They use an apt metaphor of a cake to illustrate the incompleteness of reductive models. Consider the flavor of a cake and think of the ingredients listed in the recipe. The recipe includes ingredients such as flour, sugar, shortening, vanilla extract, eggs, and milk. Does raw flour taste like cake? Does sugar, vanilla extract, or any of the other ingredients taste like cake? They do not, and knowing the individual flavors of each ingredient does not tell us much about what cake tastes like. Even mixing all of the ingredients in the correct proportions does not get us cake. Instead, external factors such as baking at the right temperature, for the right amount of time, and even the particularities of our evolved sense of taste and smell are all necessary components of experiencing the cake.<\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ff00ff\">Lewontin, Rose, and Kamin (2017) argue that the same is true for humans and other organisms.<\/span><\/p>\n<p class=\"import-Normal\">Knowing everything about cake ingredients does not allow us to fully know cake. Equally so, knowing everything about the genes found in our DNA does not allow us to fully know humans. Different, interacting levels are implicated in the development and evolution of all organisms, including humans. Genes, the structure of chromosomes, developmental processes, epigenetic tags, environmental factors, and still-other components all play key roles such that genetically reductive models of human development and evolution are woefully inadequate.<\/p>\n<p class=\"import-Normal\">The complex interactions across many levels\u2014genetic, developmental, and environmental\u2014explain why we still do not know how our one-dimensional DNA nucleotide sequence results in a four-dimensional organism. This was the unfulfilled promise of the inception of the Human Genome Project in the 1980s and 1990s: the project produced the complete DNA sequence of a human cell in the hopes that it would reveal how human bodies are built and how to cure them when they are built poorly. Yet, that information has remained elusive. Presumably, the knowledge of how organisms are produced from DNA sequences will one day permit us to reconcile the discrepancies between patterns in anatomical evolution and molecular evolution.<\/p>\n<p class=\"import-Normal\">In this chapter, we will consider multilevel evolution and explore evolution as a complex interaction between genetic and epigenetic factors as well as the environments in which organisms live. Next, we will examine the biopolitical nature of human evolution. We will then investigate problems that arise from attributing all traits to an adaptive function. Finally, we will address common misconceptions about evolution. The goal of this chapter is to provide you with the necessary toolkit for understanding the molecular, anatomical, and political dimensions of evolution.<\/p>\n<h2 class=\"import-Normal\">Evolution Happens at Multiple Levels<\/h2>\n<p class=\"import-Normal\">Following Richard Dawkins\u2019s publication of <em>The Selfish Gene <\/em>in 1976, the scientific imagination was captured by the potential of genomics to reveal how genes are copied by Darwinian selection. Dawkins argues that the genes in individuals that contribute to greater reproductive success are the units of selection. His conception of evolution at the molecular level undercuts the complex interactions between organisms and their environments, which are not expressed genomically but are nevertheless key drivers in evolution.<\/p>\n<p class=\"import-Normal\">By the 1980s, the acknowledgment among most biologists that even though genes construct bodies, genes and bodies evolve at different rates and with distinct patterns. This realization led to a renewed focus on how bodies change. The Evolutionary Synthesis of the 1930s\u20131970s had reduced organisms to their <strong>genotypes<\/strong> and species to their <strong>gene pools<\/strong>, which provided valuable insights about the processes of biological change, but it was only a first approximation. Animals are in fact reactive and adaptable beings, not passive and inert genotypes. Species are clusters of socially interacting and reproductively compatible organisms.<\/p>\n<figure style=\"width: 291px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/06\/image8-5.png\" alt=\"An asteroid hits the ocean. Pterodactyls fly among clouds in the foreground.\" width=\"291\" height=\"233\" \/><figcaption class=\"wp-caption-text\">Figure 17.1: A painting by Donald E. Davis representing the Chicxulub asteroid impact off the Yucatan Peninsula that contributed to the mass extinction that included the dinosaurs about 65 million years ago. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Chicxulub_impact_-_artist_impression.jpg\">Chicxulub impact - artist impression<\/a> by Donald E. Davis, <a href=\"https:\/\/www.nasa.gov\/\">NASA<\/a>, is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Once we accept that evolutionary change is fundamentally genetic change, we can ask: How do bodies function and evolve? How do groups of animals come to see one another as potential mates or competitors for mates, as opposed to just other creatures in the environment? Are there evolutionary processes that are not explicable by population genetics? These questions\u2014which lead us beyond reductive assumptions\u2014were raised in the 1980s by Stephen Jay Gould, the leading evolutionary biologist of the late 20th century (see: Gould 2003; 1996).<\/p>\n<p class=\"import-Normal\">Gould spearheaded a movement to identify and examine higher-order processes and features of evolution that were not adequately explained by population genetics. For example, <strong>extinction<\/strong>, which was such a problem for biologists of the 1600s, could now be seen as playing a more complex role in the history of life than population genetics had been able to model. Gould recognized that there are two kinds of extinctions, each with different consequences: background extinctions and mass extinctions. Background extinctions are those that reflect the balance of nature, because in a competitive Darwinian world, some things go extinct and other things take their place. Ecologically, your species may be adapted to its niche, but if another species comes along that\u2019s better adapted to the same niche, eventually your species will go extinct. It sucks, but it is the way of all life: you come into existence, you endure, and you pass out of existence. But mass extinctions are quite different. They reflect not so much the balance of nature as the wholesale disruption of nature: many species from many different lineages dying off at roughly the same time\u2014presumably as the result of some kind of rare ecological disaster. The situation may not be survival of the fittest as much as survival of the luckiest. The result, then, would be an ecological scramble among the survivors. Having made it through the worst, the survivors could now simply divide up the new ecosystem amongst themselves, since their competitors were gone. Something like this may well have happened about 65 million years ago, when a huge asteroid hit the Yucatan Peninsula, which mammals survived but dinosaurs did not (Figure 17.1). Something like this may be happening now, due to human expansion and environmental degradation. Note, though, that there is only a limited descriptive role here for population genetics: the phenomena we are describing are about organisms and species in ecosystems.<\/p>\n<p class=\"import-Normal\">Another question involved the disconnect between properties of <em>species<\/em> and the properties of <em>gene pools<\/em>. For example, there are upwards of 15 species of gibbons but only two species of chimpanzees. Why? There are upwards of 20 species of guenons but fewer than ten of baboons. Why? Are there genes for that? It seems unlikely. Gould suggested that species, as units of nature, might have properties that are not reducible to the genes in their cells. For example, rates of speciation and extinction might be properties of their ecologies and histories rather than their genes. Thus, relationships between environmental contexts and variability within a species result in degrees of resistance to extinction and affect the frequency and rates at which clades diversify (Lloyd and Gould 1993). Consistent biases of speciation rates might well produce patterns of macroevolutionary diversity that are difficult to explain genetically and better understood ecologically. Gould called such biases in speciation rates <strong>species selection<\/strong>\u2014a higher-order process that invokes competition between species, in addition to the classic Darwinian competition between individuals.<\/p>\n<p class=\"import-Normal\">One of Gould\u2019s most important studies involved the very nature of species. In the classical view, a species is continually adapting to its environment until it changes so much that it is a different species than it was at the beginning of this sentence (Eldredge and Gould 1972). That implies that the species is a fundamentally unstable entity through time, continuously changing to fit in. But suppose, argued Gould along with paleontologist Niles Eldredge, a species is more stable through time and only really adapts during periods of ecological instability and change. Then we might expect to find in the fossil record long equilibrium periods\u2014a few million years or so\u2014in which species don\u2019t seem to change much, punctuated by relatively brief periods in which they change a bit and then stabilize again as new species. They called this idea <strong>punctuated equilibria<\/strong>. The idea helps to explain certain features of the fossil record, notably the existence of small anatomical \u201cgaps\u201d between closely related fossil forms (Figure 17.2). Its significance lies in the fact that although it incorporates genetics, punctuated equilibria is not really a theory of genetics but one of types bodies in deep time.<\/p>\n<p class=\"import-Normal\">Punctuated equilibria is seen across taxa, with long periods in the fossil record representing little phenotypic change. These periods of stability are disrupted by shorter periods of rapid <strong>adaptation<\/strong>, the process through which populations of organisms become suited to living in their environments. Phenotypic changes are often coupled with drastic climatic or ecological changes that affect the milieu in which organisms live. For example, throughout much of hominin evolutionary history, brain size was closely associated with body size and thus remained mostly stable. However, changes occurred in average hominin brain size at around 100 thousand years ago, 1 million years ago, and 1.8 million years ago. Several hypotheses have been put forth to explain these changes, including unpredictability in climate and environment (Potts 1998), social development (Barton 1996), and the evolution of language (Deacon 1998). Evidence from the fossil record, paleoclimate models, and comparative anatomy suggests that the changes observed in hominin lineage result from biocultural processes\u2014that is, the coalescence of environmental and cultural factors that selected for larger brains (Marks 2015; Shultz, Nelson, and Dunbar 2012).<\/p>\n<figure style=\"width: 461px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image5-8.png\" alt=\"Two graphs contrast phyletic gradualism and punctuated equilibria.\" width=\"461\" height=\"222\" \/><figcaption class=\"wp-caption-text\">Figure 17.2: Different ways of conceptualizing the evolutionary relationship between an earlier and a later species. With phyletic gradualism, species are envisioned transforming continually in a direct line over time. With punctuated equilibria species branch off at particular points over time.\u00a0 Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__\/\">Phyletic gradualism vs. punctuated equilibria (Figure 2.12)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">In response to the call for a theory of the evolution of form, the field of <strong>evo-devo<\/strong>\u2014the intersection of evolutionary and developmental biology\u2014arose. The central focus here is on how changes in form and shape arise. An embryo matures by the stimulation of certain cells to divide, forming growth fields. The interactions and relationships among these growth fields generate the structures of the body. The <strong>hox genes<\/strong> that regulate these growth fields turn out to be highly conserved across the animal kingdom. This is because they repeatedly turn on and off the most basic genes guiding the animal\u2019s development, and thus any changes to them would be catastrophic. Indeed, these genes were first identified by manipulating them in fruit flies, such that one could produce a bizarre mutant fruit fly that grew a pair of legs where its antennae were supposed to be (Kaufman, Seeger, and Olsen 1990).<\/p>\n<p class=\"import-Normal\">Certain genetic changes can alter the fates of cells and the body parts, while other genetic changes can simply affect the rates at which neighboring groups of cells grow and divide, thus producing physical bumps or dents in the developing body. The result of altering the relationships among these fields of cellular proliferation in the growing embryo is <strong>allometry<\/strong>, or the differential growth of body parts. As an animal gets larger\u2014either over the course of its life or over the course of macroevolution\u2014it often has to change shape in order to live at a different size. Many important physiological functions depend on properties of geometric area: the strength of a bone, for example, is proportional to its cross-sectional area. But area is a two-dimensional quality, while growing takes place in three dimensions\u2014as an increase in mass or volume. As an animal expands, its bones necessarily weaken, because volume expands faster than area does. Consequently a bigger animal has more stress on its bones than a smaller animal does and must evolve bones even thicker than they would be by simply scaling the animal up proportionally. In other words, if you expand a mouse to the size of an elephant, it will nevertheless still have much thinner bones than the elephant does. But those giant mouse bones will unfortunately not be adequate to the task. Thus, a giant mouse would have to change aspects of its form to maintain function at a larger size (see Figure 17.3).<\/p>\n<p class=\"import-Normal\"><img class=\"aligncenter\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image6-6.png\" alt=\"Side-view of a mouse skeleton.\" width=\"515\" height=\"252\" \/><\/p>\n<figure style=\"width: 453px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image7-9.png\" alt=\"Side-view of an elephant skeleton.\" width=\"453\" height=\"326\" \/><figcaption class=\"wp-caption-text\">Figure 17.3: Mouse (top) and elephant (bottom) skeletons. Notice the elephant\u2019s bones are more robust when the two animals are the same size. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__\/\">Mouse and elephant skeletons (Figure 2.13)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Physiologically, we would like to know how the body \u201cknows\u201d when to turn on and off the genes that regulate growth to produce a normal animal. Evolutionarily, we would like to know how the body \u201clearns\u201d to alter the genetic on\/off switch (or the genetic \u201cslow down\/speed up\u201d switch) to produce an animal that looks different. Moreover, since organisms differ from one another, we would like to know how the developing body distinguishes a range of normal variation from abnormal variation. And, finally, how does abnormal variation eventually become normal in a descendant species?<\/p>\n<p class=\"import-Normal\">Taking up these questions, Gould invoked the work of a British geneticist named Conrad H. Waddington, who thought about genetics in less reductive ways than his colleagues. Rather than isolate specific DNA sites to analyze their function, Waddington instead studied the inheritance of an organism\u2019s reactivity\u2014its ability to adapt to the circumstances of its life. In a famous experiment, he grew fruit fly eggs in an atmosphere containing ether. Most died, but a few survived somehow by developing a weird physical feature: a second thorax with a second pair of wings. Waddington bred these flies and soon developed a stable line of flies who would reliably develop a second thorax when grown in ether. Then he began to lower the concentration of ether, while continuing to selectively breed the flies that developed the strange appearance. Eventually he had a line of flies that would stably develop the \u201cbithorax\u201d <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1238\"><strong>phenotype<\/strong><\/a>\u2013the suite of traits of an organism\u2013even when there was no ether; it had become the \u201cnew normal.\u201d The flies had genetically assimilated the bithorax condition.<\/p>\n<p class=\"import-Normal\">Waddington was thus able to mimic the <strong>inheritance of acquired characteristics<\/strong>: what had been a trait stimulated by ether a few generations ago was now a normal part of the development of the descendants. Waddington recognized that while he had performed a selection experiment on genetic variants, he had not selected for particular traits. Rather, he helped produce the physiological tendency to develop particular traits when appropriately stimulated. He called that tendency <strong>plasticity<\/strong> and its converse, the tendency to stay the same even under weird environmental circumstances, <strong>canalization.<\/strong> Waddington had initially selected for plasticity, the tendency to develop the bithorax phenotype under weird conditions, and then, later, for canalization, the developmental normalization of that weird physical trait. Although Waddington had high stature in the community of geneticists, evolutionary biologists of the 1950s and 1960s regarded him with suspicion because he was not working within the standard mindset of reductionism, which saw evolution as the spread of genetic variants that coded for favorable traits. Both Waddington and Gould resisted contemporary intellectual paradigms that favored reductive accounts of evolutionary processes. They conceived of evolution as an emergent process in which many external factors (e.g. climate, environment, predation) and internal factors (e.g., genotypes, plasticity, canalization) coalesce to produce the evolutionary trends that we observe in the fossil record and our genome.<\/p>\n<p class=\"import-Normal\">While Gould and Waddington both looked beyond the genome to understand evolution, the Human Genome Project\u2014an international project with the goal of identifying each base pair in the human genome in the 1990s\u2014generated a great deal of public interest in analyzing the human DNA sequence from the standpoint of medical genetics. Some of the rhetoric aimed to sell the public on investing a lot of money and resources in sequencing the human genome in order to show the genetic basis of heritable traits, cure genetic diseases, and learn what it means ultimately to be biologically human. However, the Human Genome Project was not actually able to answer those questions through the use of genetics alone, and thus a broader, more holistic account was required.<\/p>\n<p class=\"import-Normal\">This holistic account came from decades of research in human biology and anthropology, which understood the human body as highly adaptable, dynamic, and emergent. For example, in the early 20th century, anthropologist Franz Boas measured the skulls of immigrants to the U.S., revealing that environmental, not merely genetic, factors affected skull shape. The growing human body adjusts itself to the conditions of life, such as diet, sunshine, high altitude, hard labor, population density, how babies are carried\u2014any and all of which can have subtle but consistent effects upon its development. There can thus be no normal human form, only a context-specific range of human forms.<\/p>\n<p class=\"import-Normal\">However, what the human biologists called human adaptability, evolutionary biologists called developmental plasticity, and evidence quickly began to mount for its cause being <strong>epigenetic <\/strong>modifications to DNA. Epigenetic modifications are changes to how genes are used by the body (as opposed to changes in the DNA sequences; see Chapter 3). Scientific interest shifted from the focus of the Human Genome Project to the ways that bodies are made by evolutionary-developmental processes, including epigenetics. What is meant by \u201cepigenetic modification\u201d? Evolution is about how descendants diverge from their ancestors. Inheritance from parent to offspring is still critical to this process, which occurs through genetic recombination: the pairing of homologous chromosomes and sharing of genetic material during meiosis (see Chapter 3). However, in the 21st century, the link between evolution and inheritance has broadened with a clearer understanding of how environmental and developmental factors shape bodies and the expression of genes, including epigenetic inheritance patterns. While offspring inherit their genes through random assortment during meiosis, environmental factors also shape how genes are used. When these epigenetic modifications occur in germ cells, they can be passed onto offspring. In these cases, there is no change in the DNA sequence but rather in how genes are used by the body due to DNA methylation and the structure of chromosomes due to histone acetylation (see Chapter 3).<\/p>\n<p class=\"import-Normal\">In addition, we now recognize that evolution is affected by two other forms of intergenerational transmission and inheritance (in addition to genetics and epigenetics). These forms include behavioral variation and culture. That is, behavioral information can be transmitted horizontally (intragenerationally), permitting more rapid ways for organisms to adjust to the environment. And, then there is the fourth mode of transmission: the cultural or symbolic mode. <span style=\"background-color: #ffff00\">Humans are the only species<\/span> that horizontally transmits an arbitrary set of rules to govern communication, social interaction, and thought. This shared information is symbolic and has resulted in what we recognize as \u201cculture\u201d: locally emergent worlds of names, words, pictures, classifications, revered pasts, possible futures, spirits, dead ancestors, unborn descendants, in-laws, politeness, taboo, justice, beauty, and story, all accompanied by practices and a material world of tools.<\/p>\n<p class=\"import-Normal\">Consequently our contemporary ideas about evolution see the evolutionary processes as hierarchically organized and not restricted to the differential transmission of DNA sequences into the next generation. While that is indeed a significant part of evolution, the organism and species are nevertheless crucial to understanding how those DNA sequences get transmitted. Further, the transmission of epigenetic, behavioral, and symbolic information play a complex role in perpetuating our genes, bodies, and species. In the case of human evolution, one can readily see that symbolic information and cultural adaptation are far more central to our lives and our survival today than DNA and genetic adaptation. It is thus misleading to think of humans passively occupying an environmental niche. Rather, humans are actively engaged in constructing our own niches, as well as adapting to them and using them to adapt. The complex interplay between a species and its active engagement in creating its own ecology is known as <strong>niche construction<\/strong>. If we understand <strong>natural selection<\/strong>\u2013the process by which populations adapt to their specific environments\u2013as the effects that environmental context has on the reproductive success of organisms, then niche construction is the process through which organisms shape their own selective pressures.<\/p>\n<h2 class=\"import-Normal\">The Biopolitics of Heredity<\/h2>\n<p class=\"import-Normal\">\u201cScience isn\u2019t political\u201d is a sentiment that you have likely heard before. Science is supposed to be about facts and objectivity. It exists, or at least ought to, outside of petty human concerns. However, the sorts of questions we ask as scientists, the problems we choose to study, the categories and concepts we use, who gets to do science, and whose work gets cited are all shaped by culture. Doing science is a political act. This fact is markedly true for human evolution. While it is easier to create intellectual distance between us and fruit flies and viruses, there is no distance when we are studying ourselves. The hardest lesson to learn about human evolution is that it is intensely political. Indeed, to see it from the opposite side, as it were, the history of creationism\u2014the belief that the universe was divinely created around 6,000 years ago\u2014is essentially a history of legal decisions. For instance, in <em>Tennessee v. John T. Scopes<\/em> (1925), a schoolteacher was prosecuted for violating a law in Tennessee that prohibited the teaching of human evolution in public schools, where teachers were required by law to teach creationism.<\/p>\n<p class=\"import-Normal\">More recently, legal decisions aimed at legislating science education have shaped how students are exposed to evolutionary theory. For instance, <em>McLean v. Arkansas<\/em> (1982) dispatched \u201cscientific creationism\u201d by arguing that the imposition of balanced teaching of evolution and creationism in science classes violates the Establishment Clause, separating church and state. Additionally, <em>Kitzmiller v. Dover (Pennsylvania) Area School District<\/em> (2005) dispatched the teaching of \u201cintelligent design\u201d in public school classrooms as it was deemed to not be science. In some cases, people see unbiblical things in evolution, although most Christian theologians are easily able to reconcile science to the Bible. In other cases, people see immoral things in evolution, although there is morality and immorality everywhere. And some people see evolution as an aspect of alt-religion, usurping the authority of science in schools to teach the rejection of the Christian faith, which would be unconstitutional due to the protected separation of church and state.<\/p>\n<p class=\"import-Normal\">Clearly, the position that politics has nothing to do with science is untenable. But is the politics in evolution an aberration or is it somehow embedded in science? In the early 20th century, scientists commonly promoted the view that science and politics were separate: science was seen as a pure activity, only rarely corrupted by politics. And yet as early as World War I, the politics of nationalism made a hero of the German chemist Fritz Haber for inventing poison gas. And during World War II, both German doctors and American physicists, recruited to the war effort, helped to end many civilian lives. Therefore, we can think of the apolitical scientist as a self-serving myth that functions to absolve scientists of responsibility for their politics. The history of science shows how every generation of scientists has used evolutionary theory to rationalize political and moral positions. In the very first generation of evolutionary science, Darwin\u2019s <em>Origin of Species<\/em> (1859) is today far more readable than his <em>Descent of Man<\/em> (1871). The reason is that Darwin consciously purged <em>The Origin of Species<\/em> of any discussion of people. And when he finally got around to talking about people, in <em>The Descent of Man<\/em>, he simply imbued them with the quaint Victorian prejudices of his age, and the result makes you cringe every few pages. There is plenty of politics in there\u2014sexism, racism, and colonialism\u2014because <em>you cannot talk about people apolitically<\/em>.<\/p>\n<p class=\"import-Normal\">One immediate faddish deduction from Darwinism, popularized by Herbert Spencer (1864) as \u201csurvival of the fittest,\u201d held that unfettered competition led to advancement in nature and to human history. Since the poor were purported losers in that struggle, anything that made their lives easier would go against the natural order. This position later came to be known ironically as \u201cSocial Darwinism.\u201d Spencer was challenged by fellow Darwinian Thomas Huxley (1863), who agreed that struggle was the law of the jungle but observed that we don\u2019t live in jungles anymore. The obligation to make lives better for others is a moral, not a natural, fact. We simultaneously inhabit a natural universe of descent from apes and a moral universe of injustice and inequality, and science is not well served by ignoring the latter.<\/p>\n<p class=\"import-Normal\">Concurrently, the German biologist Ernst Haeckel\u2019s 1868 popularization of Darwinism was translated into English a few years later as <em>The History of Creation<\/em>. As we saw earlier, Haeckel was determined to convince his readers that they were descended from apes, even in the absence of fossil evidence attesting to it. When he made non-Europeans into the missing links that connected his readers to the apes, and depicted them as ugly caricatures, he knew precisely what he was doing. Indeed, even when the degrading racial drawings were deleted from the English translation of his book, the text nevertheless made his arguments quite clear. And a generation later, when the Americans had not yet entered the Great War in 1916, a biologist named Vernon Kellogg visited the German High Command as a neutral observer and found that the officers knew a lot about evolutionary biology, which they had gotten from Haeckel and which rationalized their military aggressions. Kellogg went home and wrote a bestseller about it, called <em>Headquarters Nights<\/em> (1917). World War I would have been fought with or without evolutionary theory, but as a source of scientific authority, evolution\u2014even if a perversion of the Darwinian theory\u2014had very quickly attained global geopolitical relevance.<\/p>\n<p class=\"import-Normal\">Oftentimes, politics in evolutionary science is subtle, due to the pervasive belief in the advancement of science. We recognize the biases of our academic ancestors and modify our scientific stories accordingly. But we can never be free of our own cultural biases, which are invisible to us, as much as our predecessors\u2019 biases were invisible to them. In some cases, the most important cultural issues resurface in different guises each generation, like scientific racism. <strong>Scientific racism<\/strong> is the recruitment of science for the evil political ends of racism, and it has proved remarkably impervious to evolution. Before Darwin, there was creationist scientific racism, and after Darwin, there was evolutionist scientific racism. And there is still scientific racism today, self-justified by recourse to evolution, which means that scientists have to be politically astute and sensitive to the uses of their work to counter these social tendencies.<\/p>\n<p class=\"import-Normal\">Consider this: Are you just your ancestry, or can you transcend it? If that sounds like a weird question, it was actually quite important to a turn-of-the-20th-century European society in which an old hereditary aristocracy was under increasing threat from a rising middle class. And that is why the very first English textbook of Mendelian genetics concluded with the thought that \u201cpermanent progress is a question of breeding rather than of pedagogics; a matter of gametes, not of training \u2026 the creature is not made but born\u201d (Punnett 1905, 60). <em>Translation: Not only do we now know a bit about how heredity works, but it\u2019s also the most important thing about you. Trust me, I\u2019m a scientist.<\/em><\/p>\n<p class=\"import-Normal\">Yet evolution is about how descendants come to differ from ancestors. Do we really know that your heredity, your DNA, your ancestry, is the most important thing about you? That you were born, not made? After all, we do know that you could be born into slavery or as a peasant, and come from a long line of enslaved people or peasants, and yet not have slavery or peasantry be the most important thing about you. Whatever your ancestors were may unfortunately constrain what you can become, but as a moral precept, it should not. But just as science is not purely \u201cfacts and objectivity,\u201d ancestry is not a strictly biological concept. Human ancestry is biopolitics, not biology.<\/p>\n<p class=\"import-Normal\">Evolution is fundamentally a theory about ancestry, and yet ancestors are, in the broad anthropological sense, sacred: ancestors are often more meaningful symbolically than biologically. Just a few years after <em>The Origin of Species <\/em>(Darwin 1859), the British politician and writer Benjamin Disraeli declared himself to be on the side of the angels, not the apes, and to \u201crepudiate with indignation and abhorrence those new-fangled theories\u201d (Monypenny, Flavelle, and Buckle 1920, 105). He turned his back on an ape ancestry and looked to the angel; yet, he did so as a prominent Jew-turned-Anglican, who had personally transcended his humble roots and risen to the pinnacle of the Empire. Ancestry was certainly important, and Disraeli was famously proud of his, but it was also certainly not the most important thing, not the primary determinant of his place in the world. Indeed, quite the opposite: Disraeli\u2019s life was built on the transcendence of many centuries of Jewish poverty and oppression in Europe. Humble ancestry was there to be superseded and nobility was there to be earned; Disraeli would later become the Earl of Beaconsfield. Clearly, \u201care you just your ancestry\u201d is not a value-neutral question, and \u201cthe creature is not made, but born\u201d is not a value-neutral answer.<\/p>\n<p class=\"import-Normal\">Ancestry being the most important thing about a person became a popular idea twice in 20th century science. First, at the beginning of the century, when the <strong>eugenics<\/strong> movement in America called attention to \u201cfeeble-minded stocks,\u201d which usually referred to the poor or to immigrants (see Figure 17.4; and see Chapter 2). This movement culminated in Congress restricting the immigration of \u201cfeeble-minded races\u201d (said to include Jews and Italians) in 1924, and the Supreme Court declaring it acceptable for states to sterilize their \u201cfeeble-minded\u201d citizens involuntarily in 1927. After the Nazis picked up and embellished these ideas during World War II, Americans moved swiftly away from them in some contexts (e.g., for most people of European descent) while still strictly adhering in other contexts (e.g., Japanese internment camps and immigration restrictions).<\/p>\n<figure style=\"width: 374px\" class=\"wp-caption alignright\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image4-6.png\" alt=\"Historic photo. People sit in front of a structure with a \u201cEugenic and Health Exhibit&quot; banner.\" width=\"374\" height=\"262\" \/><figcaption class=\"wp-caption-text\">Figure 17.4: Eugenic and Health Exhibit, Fitter Families exhibit, and examination building, Kansas State Free Fair. Credit: <a href=\"https:\/\/www.dnalc.org\/view\/16328-Gallery-14-Eugenics-Exhibit-at-the-Kansas-State-Free-Fair-1920.html\">Gallery 14: Eugenics Exhibit at the Kansas State Free Fair, 1920 ID (ID 16328)<\/a> by <a href=\"https:\/\/www.dnalc.org\/\">Cold Spring Harbor<\/a> (Courtesy American Philosophical Society) is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/3.0\/us\/\">CC BY-NC-ND 3.0 License.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\">Ancestry again became paramount in the drumming up of public support for the Human Genome Project in the 1990s. Public support for sequencing the human genome was encouraged by a popular science campaign that featured books titled <em>The Book of Man <\/em>(Bodmer and McKie 1997), <em>The Human Blueprint <\/em>(Shapiro 1991), and <em>The Code of Codes<\/em> (Kevles and Hood 1993). These books generally promised cures for genetic diseases and a deeper understanding of the human condition. We can certainly identify progress in molecular genetics over the last couple of decades since the human genome was sequenced, but that progress has notably not been accompanied by cures for genetic diseases, nor by deeper understandings of the human condition.<\/p>\n<p class=\"import-Normal\">Even at the most detailed and refined levels of genetic analysis, we still don\u2019t have much of an understanding of the actual basis by which things seem to \u201crun in families.\u201d While the genetic basis of simple, if tragic, genetic diseases have become well-known\u2014such as sickle-cell anemia, cystic fibrosis, and Tay-Sachs\u2019 Disease\u2014we still haven\u2019t found the ostensible genetic basis for traits that are thought to have a strong genetic component. For example, a recent genetic summary found over 12,000 genetic sites that contributed to height yet still explained only about 40-50 percent of the variation in height among European ancestry but no more than 10-20 percent of variation of other ancestries, which we know strongly runs in families (Yengo et al. 2022).<\/p>\n<p class=\"import-Normal\">Partly in reaction to the reductionistic hype of the Human Genome Project, the study of epigenetics has become the subject of great interest. One famous natural experiment involves a Nazi-imposed famine in Holland over the winter of 1944\u20131945. Children born during and shortly after the famine experienced a higher incidence of certain health problems as adults, many decades later. Apparently, certain genes had been down-regulated early in development and remained that way throughout the course of life. Indeed, this modified regulation of the genes in response to the severe environmental conditions may have been passed on to their children.<\/p>\n<p class=\"import-Normal\">Obviously one\u2019s particular genetic constitution may play an important role in one\u2019s life trajectory. But overvaluing that role may have important social and political consequences. In the first place, genotypes are rendered meaningful in a cultural universe. Thus, if you live in a strongly patriarchal society and are born without a Y chromosome (since human males are chromosomally XY and females XX), your genotype will indeed have a strong effect upon your life course. So even though the variation is natural, the consequences are political. The mediating factors are the cultural ideas about how people of different sexes ought to be treated, and the role of the state in permitting certain people to develop and thrive. More broadly, there are implications for public education if variation in intelligence is genetic. There are implications for the legal system if criminality is genetic. There are implications for the justice system if sexual preference, or sexual identity, is genetic. There are implications for the development of sports talent if that is genetic. And yet, even for the human traits that are more straightforward to measure and known to be strongly heritable, the DNA base sequence variation seems to explain little.<\/p>\n<p class=\"import-Normal\">Genetic determinism or <strong>hereditarianism<\/strong> is the idea that \u201cthe creature is made, not born\u201d\u2014or, in a more recent formulation by James Watson, that \u201cour fate is in our genes.\u201d One of the major implications drawn from genetic determinism is that the feature in question must inevitably express itself; therefore, we can\u2019t do anything about it. Therefore, we might as well not fund the social programs designed to ameliorate economic inequality and improve people\u2019s lives, because their courses are fated genetically. And therefore, they don\u2019t deserve better lives.<\/p>\n<p class=\"import-Normal\">All of the \u201ctherefores\u201d in the preceding paragraph are open to debate. What is important is that the argument relies on a very narrow understanding of the role of genetics in human life, and it misdirects the causes of inequality from cultural to natural processes. By contrast, instead of focusing on genes and imagining them to place an invisible limit upon social progress, we can study the ways in which your DNA sequence does <em>not<\/em> limit your capability for self-improvement or fix your place in a social hierarchy. In general, two such avenues exist. First, we can examine the ways in which the human body responds and reacts to environmental variation: human adaptability and plasticity. This line of research began with the anthropometric studies of immigrants by Franz Boas in the early 20th century and has now expanded to incorporate the epigenetic inheritance of modified human DNA. And second, we can consider how human lives are shaped by social histories\u2014especially the structural inequalities within the societies in which they grow up.<\/p>\n<p class=\"import-Normal\">Although it arises and is refuted every generation, the radical hereditarian position (genetic determinism) perennially claims to speak for both science and evolution. It does not. It is the voice of a radical fringe\u2014perhaps naive, perhaps evil. It is not the authentic voice of science or of evolution. Indeed, keeping Charles Darwin\u2019s name unsullied by protecting it from association with bad science often seems like a full-time job. Culture and epigenetics are very much a part of the human condition, and their roles are significant parts of the complete story of human evolution.<\/p>\n<p><span style=\"background-color: #00ffff\"><span style=\"text-decoration: underline\">(Sterilization of Indigenous women in Canada)<\/span> (https:\/\/www.thecanadianencyclopedia.ca\/en\/article\/sterilization-of-indigenous-women-in-canada)\u00a0<\/span><\/p>\n<h2 class=\"import-Normal\">Adaptationism and the Panglossian Paradigm<\/h2>\n<p class=\"import-Normal\">The story of human evolution, and the evolution of all life for that matter, is never settled because evolution is ongoing. Additionally, because the conditions that shape evolutionary trajectories are not predetermined, evolution itself is emergent. Even during periods of ecological stability, when fewer macroevolutionary changes occur, populations of organisms continue to experience change. When ecological stability is disrupted, populations must adapt to the changes. Darwin explained in naturalistic terms how animals adapt to their environments: traits that contribute to an organism's ability to survive and reproduce in specific environments will become more common. The most \u201cfit\u201d\u2014those organisms best suited to the <em>current<\/em> environmental conditions in which they live\u2014have survived over eons of the history of life on earth to cocreate ecosystems full of animals and plants. Our own bodies are full of evident adaptations: eyes for seeing, ears for hearing, feet for walking on, and so forth.<\/p>\n<p class=\"import-Normal\">But what about hands? Feet are adapted to be primarily weight-bearing structures (rather than grasping structures, as in the apes) and that is what we primarily use them for. But we use our hands in many ways: for fine-scale manipulation, greeting, pointing, stimulating a sexual partner, writing, throwing, and cooking, among other uses. So which of these uses express what hands are \u201cfor,\u201d when all of them express what hands do?<\/p>\n<p class=\"import-Normal\">Gould and Lewontin (1979) illustrate the problem with assuming that the function of a trait defines its evolutionary cause. Consider the case of Dr. Pangloss\u2014the protagonistic of Voltaire\u2019s <em>Candide<\/em>\u2014who believed that we lived in the best of all possible worlds. Gould and Lewontin use his pronouncement that \u201cnoses were made for spectacles and so we have spectacles\u201d to demonstrate the problem with assuming any trait has evolved for a specific purpose. Identifying a function of a trait does not necessitate that the function is the ultimate cause of the trait. Individual traits are not under selection pressures in isolation; in fact, an entire organism must be able to survive and reproduce in their environment. When natural selection results in adaptations, changes that occur in some traits can have cascading effects throughout the phenotype and features that are not under selection pressure can also change.<\/p>\n<figure style=\"width: 279px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image3-5.png\" alt=\"Human hand is smaller with smaller fingers and smoother skin compared to a chimpanzee hand.\" width=\"279\" height=\"264\" \/><figcaption class=\"wp-caption-text\">Figure 17.5: Drawings of a human hand (left) and a chimpanzee hand (right). Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__\/\">Human and chimpanzee hand (Figure 2.16)<\/a> by Mary Nelson original to <a href=\"https:\/\/explorations.americananthro.org\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">There is an important lesson in recognizing that what things do in the present is not a good guide to understanding why they came to exist. Gunpowder was invented for entertainment\u2014only later was it adopted for killing people. The Internet was invented to decentralize computers in case of a nuclear attack\u2014and only later adopted for social media. Apes have short thumbs and use their hands in locomotion; our ancestors stopped using their hands in locomotion by about six million years ago and had fairly modern-looking hands by about two million years ago. We can speculate that a combination of selection for abstract thought and dexterity led to evolution of the human hand, with its capability for toolmaking that exceeds what apes can do (see Figure 17.5). But let\u2019s face it\u2014how many tools have you made today?<\/p>\n<p class=\"import-Normal\">Consequently, we are obliged to see the human foot as having a purpose to which it is adapted and the human hand as having multiple purposes, most of which are different from what it originally evolved for. Paleontologists Gould and Elisabeth Vrba suggested that an original use be regarded as an adaptation and any additional uses be called \u201c<strong>exaptations.<\/strong>\u201d Thus, we would consider the human hand to be an adaptation for toolmaking and an exaptation for writing. So how do we know whether any particular feature is an adaptation, like the walking foot, rather than an exaptation, like the writing hand? Or more broadly, how can we reason rigorously from what a feature does to what it evolved for?<\/p>\n<p class=\"import-Normal\">The answer to the question \u201cwhat did this feature evolve for?\u201d creates an origin myth. This origin myth contains three assumptions: (1) features can be isolated as evolutionary units; (2) there is a specific reason for the existence of any particular feature; and (3) there is a clear and simplistic explanation for why the feature evolved.<\/p>\n<figure style=\"width: 378px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image9-8.png\" alt=\"Head with images and human qualities drawn on it. Journal title printed at the bottom.\" width=\"378\" height=\"437\" \/><figcaption class=\"wp-caption-text\">Figure 17.6: According to the early 19th century science of phrenology, units of personality could be mapped onto units in the head, as shown on this cover of The Phrenology Journal. Credit: <a href=\"https:\/\/wellcomecollection.org\/works\/b6skynug\">Phrenology; Chart<\/a> [slide number 5278, photo number: L0000992, original print from Dr. E. Clark, The Phrenological Journal (Know Thyself)] by <a href=\"https:\/\/wellcomecollection.org\/\">Wellcome Collection<\/a>, is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/legalcode\">CC BY 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">The first assumption was appreciated a century ago as the \u201cunit-character problem.\u201d Are the units by which the body grows and evolves the same as units we name? This is clearly not the case: we have genes and we have noses, and we have genes that affect noses, but we don\u2019t have \u201cnose genes.\u201d What is the relationship between the evolving elements that we see, identify, and name, and the elements that biologically exist and evolve? It is hard to know, but we can use the history of science as a guide to see how that fallacy has been used by earlier generations. Back in the 19th century, the early anatomists argued that since the brain contained the mind, they could map different mental states (acquisitiveness, punctuality, sensitivity) onto parts of the brain. Someone who was very introspective, say, would have an enlarged introspection part of the brain, a cranial bulge to represent the hyperactivity of this mental state. The anatomical science was known as <strong>phrenology<\/strong>, and it was predicated on the false assumption that units of thought or personality or behavior could be mapped to distinct parts of the brain and physically observed (see Figure17.6). This is the fallacy of reification, imagining that something named is something real.<\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Long alt text: Side view of human head. At the top are the words \u201cKnow Thyself.\u201d On the upper head are small illustrations and word qualities such as \u201cfriendship,\u201d \u201cself-esteem,\u201d and \u201csecretiveness.\u201d On the lower part of the man\u2019s man\u2019s face are the words <em>The Phrenological Journal and Science of Health, A First Class Monthly<\/em>. The caption at the bottom reads: \u201cSpecially devoted to the \u2018.\u2019 Contains PHRENOLOGY and PHYSIOGNOMY, with all the SIGNS OF CHARACTER, and how to read them; ETHNOLOGY, or the Natural History of Man in all his relations.\u201d (All emphases in original.)<\/span><\/p>\n<figure style=\"width: 295px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image1-8-1.png\" alt=\"A black-and-white drawing of a chimpanzee head and face.\" width=\"295\" height=\"236\" \/><figcaption class=\"wp-caption-text\">Figure 17.7: Chimpanzees have big ears. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Chimpanzee_head_sketch.png\">Chimpanzee head sketch<\/a> by <a href=\"https:\/\/de.wikipedia.org\/wiki\/Benutzer:Roger_Zenner\">Roger Zenner<\/a>, original by Brehms Tierleben (1887), is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">The second assumption, that everything has a reason, has long been recognized as a core belief of religion. Our desire to impose order and simplicity on the workings of the universe, however, does not constrain it to obey simple and orderly causes. Magic, witchcraft, spirits, and divine agency are all powerful explanations for why things happen. Consequently, it is probably not a good idea to lump natural selection in with those. Sometimes things do happen for a reason, of course, but other times things happen as byproducts of other things, or for very complicated and entangled reasons, or for no reason at all. What phenomena have reasons and thereby merit explanation? Chimpanzees have very large testicles, and we think we know why: their promiscuous sexual behavior triggers intense competition for high sperm count. But chimpanzees also have very large ears, but much less scientific attention has been paid to this trait (see Figure 17.7). Why not? Why should there be a reason for chimp testicles but not for chimp ears? What determines the kinds of features that we try to explain, as opposed to the ones that we do not? Again, the assumption that any specific feature has a reason is metaphysical; that is to say, it may be true in any particular case, but to assume it in all cases is gratuitous.<\/p>\n<p class=\"import-Normal\">And third, the possibility of knowing what the reason for any particular feature is, assuming that it has one, is a challenge for evolutionary epistemology (the theory of how we know things). Consider the big adaptations of our lineage: bipedalism and language. Nobody doubts that they are good, and they evolved by natural selection, and we know how they work. But why did they evolve? If talking and walking are simply better than not talking and not walking, then why did they evolve in just a single branch of the ape lineage in the primate family tree? We don\u2019t know what bipedalism evolved for, although there are plenty of speculations: walking long distances, running long distances, cooling the head, seeing over tall grass, carrying babies, carrying food, wading, threatening, counting calories, sexual display, and so on. Neither do we know what language evolved for, although there are speculations: coordinating hunting, gossiping, manipulating others. But it is also possible that bipedality is simply the way that a small arboreal ape travels on the ground, if it isn\u2019t in the treetops. Or that language is simply the way that a primate with small canine teeth and certain mental propensities comes to communicate. If that were true, then there might be no reason for bipedality or language: having the unique suite of preconditions and a fortuitous set of circumstances simply set them in motion, and natural selection elaborated and explored their potentials. It is possible that walking and talking simply solved problems that no other lineage had ever solved; but even if so, the fact remains that the rest of the species in the history of life have done pretty well without having solved them.<\/p>\n<p class=\"import-Normal\">It is certainly very optimistic to think that all three assumptions (that organisms can be meaningfully atomized, that everything has a reason, and that we can know the reason) would be simultaneously in effect. Indeed, just as there are many ways of adapting (genetically, epigenetically, behaviorally, culturally), there are also many ways of being nonadaptive, which would imply that there is no reason at all for the feature in question.<\/p>\n<p class=\"import-Normal\">First, there is the element of randomness of population histories. There are more cases of sickle-cell anemia among sub-Saharan Africans than other peoples, and there is a reason for it: carriers of sickle-cell anemia have a resistance to malaria, which is more frequent in parts of Africa (as discussed in Chapters 4 and 14). But there are more cases of a blood disease called variegated porphyria, a rare genetic metabolic disorder, in the Afrikaners of South Africa (descendants of mostly Dutch settlers in the 17th century) than in other peoples, and there is no reason for it. Yet we know the cause: One of the founding Dutch colonial settlers had the <strong>allele<\/strong>\u2013a variant of a gene\u2013and everyone in South Africa with it today is her descendant. But that is not a reason\u2014that is simply an accident of history.<\/p>\n<p class=\"import-Normal\">Second, there is the potential mismatch between the past and the present. The value of a particular feature in the past may be changed as the environmental circumstances change. Our species is diurnal, and our ancestors were diurnal. But beginning around a few hundred thousand years ago, our ancestors could build fires, which extended the light period, which was subsequently further amplified by lamps and candles. And over the course of the 20th century, electrical power has made it possible for people to stay up very late when it is dark\u2014working, partying, worrying\u2014to a greater extent than any other closely related species. In other words, we evolved to be diurnal, yet we are now far more nocturnal than any of our recent ancestors or close relatives. Are we adapting to nocturnality? If so, why? Does it even make any sense to speak of the human occupation of a nocturnal ape niche, despite the fact that we empirically seem to be doing just that? And if so, does it make sense to ask what the reason for it is?<\/p>\n<p class=\"import-Normal\">Third, there is a genetic phenomenon known as a selective sweep, or the hitchhiker effect. Imagine three genes\u2014A, B, and C\u2014located very closely together on a chromosome. They each have several variants, or alleles, in the population. Now, for whatever reason, it becomes beneficial to have one of the B alleles, say B4; this B4 allele is now under strong positive selection. Obviously, we will expect future generations to be characterized by mostly B4. But what was B4 attached to? Because whatever A and C alleles were adjacent to it will also be quickly spread, simply by virtue of the selection for B4. Even if the A and C alleles are not very good, they will spread because of the good B4 allele between them. Eventually the linkage groups will break up because of genetic crossing-over in future generations. But in the meantime, some random version of genes A and C are proliferating in the species simply because they are joined to superior allele B4. And clearly, the A and C alleles are there because of selection\u2014but not because of selection <em>for<\/em> them!<\/p>\n<p class=\"import-Normal\">Fourth, some features are simply consequences of other properties rather than adaptations to external conditions. We already noted the phenomenon of allometric growth, in which some physical features have to outgrow others to maintain function at an increased size. Can we ask the reason for the massive brow ridges of <em>Homo erectus<\/em>, or are brow ridges simply what you get when you have a conjunction of thick skull bones, a large face, and a sloping forehead\u2014and, thus, again would have a cause but no reason?<\/p>\n<p class=\"import-Normal\">Fifth, some features may be underutilized and on the way out. What is the reason for our two outer toes? They aren\u2019t propulsive, they don\u2019t do anything, and sometimes they\u2019re just in the way. Obviously they are there because we are descended from ancestors with five digits on their hands and feet. Is it possible that a million years from now, we will just have our three largest toes, just as the ancestors of the horse lost their digits in favor of a single hoof per limb? Or will our outer toes find another use, such as stabilizing the landings in our personal jet-packs? For the time being, we can just recognize vestigiality as another nonadaptive explanation for the presence of a given feature.<\/p>\n<p class=\"import-Normal\">Finally, Darwin himself recognized that many obvious features do not help an animal survive. Some things may instead help an animal breed. The peacock\u2019s tail feathers do not help it eat, but they do help it mate. There is competition, but only against half of the species. Darwin called this <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1084\">sexual selection<\/a><\/strong>. Its result is not a fit to the environment but, rather, a fit to the opposite sex. In some species, that is literally the case, as the male and female genitalia have specific ways of anatomically fitting together. The specific form is less important than the specific match, so inquiring about the reason for a particular form of the reproductive anatomy may be misleading. The specific form may be effectively random, as long as it fits the opposite sex and is different from the anatomies of other species. Nor is sexual selection the only form of selection that can affect the body differently from natural selection. Competition might also take place between biological units other than organisms\u2014perhaps genes, perhaps cells, or populations, or species. The spread of cultural things, such as head-binding or cheap refined fructose or forced labor, can have significant effects upon bodies, which are also not adaptations produced by natural selection. They are often adaptive physiological responses to stresses but not the products of natural selection.<\/p>\n<p class=\"import-Normal\">With so many paths available by which a physical feature might have organically arisen without having been the object of natural selection, it is unwise to assume that any individual trait is an adaptation. And that generalization applies to the best-known, best-studied, and most materially based evolutionary adaptations of our lineage. But our cultural behaviors are also highly adaptive, so what about our most familiar social behaviors? Patriarchy, hierarchy, warfare\u2014are these adaptations? Do they have reasons? Are they good for something?<\/p>\n<p class=\"import-Normal\">This is where some sloppy thinking has been troublesome. What would it mean to say that patriarchy evolved by natural selection in the human species? If, on the one hand, it means that the human mind evolved by natural selection to be able to create and survive in many different kinds of social and political regimes, of which patriarchy is one, then biological anthropologists will readily agree. If, on the other hand, it means that patriarchy evolved by natural selection, that implies that patriarchy is genetically determined (since natural selection is a genetic process) and out-reproduced the alleles for other, more egalitarian, social forms. This in turn would imply that patriarchy is an adaptation and therefore of some beneficial value in the past and has become an ingrained part of human nature today. This would be bad news, say, if you harbored ambitions of dismantling it. Dismantling patriarchy in that case would be to go against nature, a futile gesture. In other words, this latter interpretation would be a naturalistic manifesto for a conservative political platform: don\u2019t try to dismantle the patriarchy, because it is within us, the product of evolution\u2014suck it up and live with it.<\/p>\n<p class=\"import-Normal\">Here, evolution is being used as a political instrument for transforming the human genome into an imaginary glass ceiling against equality. There is thus a convergence between the pseudo-biology of crude <strong>adaptationism <\/strong>(the idea that everything is the product of natural selection) and the pseudo-biology of hereditarianism. Naturalizing inequality is not the business of evolutionary theory, and it represents a difficult moral position for a scientist to adopt, as well as a poor scientific position.<\/p>\n<div class=\"textbox shaded\">\n<p>Dig Deeper: Evolution of the Anthropocene (to be reviewed)<\/p>\n<p>As humans have caused the emergence of the Anthropocene, it is important to inform scholars about the effect of our social and cultural evolution on the rest of the world. Richard Robbins\u2019 <em>Global Problems and Culture of Capitalism<\/em> explains how the modern culture of consumption has been extremely successful at accommodating populations of people far larger than previously possible. Robbins claims that the globalization attributed to capitalism has allowed the world to make full use of its environmental resources, providing necessities and innovative technologies to humans all over the world (Robbins &amp; Dowty, 2019). In other words, capitalism is an anthropocentric cultural system that highly benefits humans and facilitates our survival with little regard to the development and survival of other forms of life. It would be of high relevance to introduce the idea that our cultural evolution and capacity to modify the environment to meet our needs have established new environmental conditions in which the human species' survival and reproduction rate expand at the detriment of ecosystems and endangerment of other primates and non-human species.<\/p>\n<p>According to a 2019 UN report, following the 16th century, the world has entered a period of extreme environmental destruction that is generating ecological modifications and has led to the extinction of at least 680 vertebrate species and over 9 percent of the domesticated mammals used for food and agriculture (United Nations, 2019). Human lifestyles are causing changes that\u2014if not taken into consideration\u2014could lead to our extinction as a species. The recognition that our evolutionary behavioural development is causing environmental destruction may be the first step for our species to take accountability for the damage that it is causing to others and prevent further damage.<\/p>\n<\/div>\n<p>&nbsp;<\/p>\n<h2 class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Concluding Thoughts<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Now that you have finished reading this chapter, you are equipped to understand the historical and political dimensions of evolution. Evolution is an ongoing process of change and diversification. Evolutionary theory is a tool that we use to understand this process. The development of evolutionary theory is shaped both by scientific innovation and political engagement. Since Darwin first articulated natural selection as an observable mechanism by which species adapt to their environments, our understanding of evolution has grown. Initially, scientists focused on the adaptive aspects of evolution. However, with the emergence of genetics, our understanding of heredity and the level at which evolution acts has changed. Genetics led to a focus on the molecular dimensions of evolution. For some, this focus resulted in reductive accounts of evolution. Further developments in our understanding of evolution shifted our view to epigenetic processes and how organisms shape their own evolutionary pressures (e.g., niche construction).<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Evolutionary theory will continue to develop in the future as we invent new technologies, describe new dimensions of biology, and experience cultural changes. Current innovations in evolutionary theory are asking us to consider evolutionary forces beyond natural selection and genetics to include the ways organisms shape their environments (niche construction), inheritances beyond genetics (inclusive inheritance), constraints on evolutionary change (developmental bias), and the ability of bodies to change in response to external factors (plasticity). The future of evolutionary theory looks bright as we continue to explore these and other dimensions. Biological anthropology is well-positioned to be a lively part of this conversation, as it extends standard evolutionary theory by considering the role of culture, social learning, and human intentionality in shaping the evolutionary trajectories of humans (Zeder 2018). Remember, at root, human evolutionary theory consists of two propositions: (1) the human species is descended from other similar species and (2) natural selection has been the primary agent of biological adaptation. Pretty much everything else is subject to some degree of contestation.<\/span><\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Review Questions<\/h2>\n<ul>\n<li class=\"import-Normal\">How is the study of your ancestors biopolitical, not just biological? Does that make it less scientific or differently scientific?<\/li>\n<li class=\"import-Normal\">What was gained by reducing organisms to genotypes and species to gene pools? What is gained by reintroducing bodies and species into evolutionary studies?<\/li>\n<li class=\"import-Normal\">How do genetic or molecular studies complement anatomical studies of evolution?<\/li>\n<li class=\"import-Normal\">How are you reducible to your ancestry? If you could meet your ancestors from the year 1700 (and you would have well over a thousand of them!), would their lives be meaningfully similar to yours? Would you even be able to communicate with them?<\/li>\n<li class=\"import-Normal\">The molecular biologist Fran\u00e7ois Jacob argued that evolution is more like a tinkerer than an engineer. In what ways do we seem like precisely engineered machinery, and in what ways do we seem like jerry-rigged or improvised contraptions?<\/li>\n<li class=\"import-Normal\">How might biological anthropology contribute to future developments in evolutionary theory?<\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\">Key Terms<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\"><strong>Adaptation<\/strong>: A fit between the organism and environment.<\/p>\n<p class=\"import-Normal\"><strong>Adaptationism<\/strong>: The idea that everything is the product of natural selection.<\/p>\n<p class=\"import-Normal\"><strong>Allele<\/strong>: A genetic variant.<\/p>\n<p class=\"import-Normal\"><strong>Allometry<\/strong>: The differential growth of body parts.<\/p>\n<p class=\"import-Normal\"><strong>Canalization<\/strong>: The tendency of a growing organism to be buffered toward normal development.<\/p>\n<p class=\"import-Normal\"><strong>Epigenetics<\/strong>: The study of how genetically identical cells and organisms (with the same DNA base sequence) can nevertheless differ in stably inherited ways.<\/p>\n<p class=\"import-Normal\"><strong>Eugenics<\/strong>: An idea that was popular in the 1920s that society should be improved by breeding \u201cbetter\u201d kinds of people.<\/p>\n<p class=\"import-Normal\"><strong>Evo-devo<\/strong>: The study of the origin of form; a contraction of \u201cevolutionary developmental biology.\u201d<\/p>\n<p class=\"import-Normal\"><strong>Exaptation<\/strong>: An additional beneficial use for a biological feature.<\/p>\n<p class=\"import-Normal\"><strong>Extinction<\/strong>: The loss of a species from the face of the earth.<\/p>\n<p class=\"import-Normal\"><strong>Gene<\/strong>: A stretch of DNA with an identifiable function (sometimes broadened to include any DNA with recognizable structural features as well).<\/p>\n<p class=\"import-Normal\"><strong>Gene pool<\/strong>: Hypothetical summation of the entire genetic composition of population or species.<\/p>\n<p class=\"import-Normal\"><strong>Genotype<\/strong>: Genetic constitution of an individual organism.<\/p>\n<p class=\"import-Normal\"><strong>Hereditarianism<\/strong>: The idea that genes or ancestry is the most crucial or salient element in a human life. Generally associated with an argument for natural inequality on pseudo-genetic grounds.<\/p>\n<p class=\"import-Normal\"><strong>Hox genes<\/strong>: A group of related genes that control for the body plan of an embryo along the head-tail axis.<\/p>\n<p class=\"import-Normal\"><strong>Inheritance of acquired characteristics<\/strong>: The idea that you pass on the features that developed during your lifetime, not just your genes; also known as Lamarckian inheritance.<\/p>\n<p class=\"import-Normal\"><strong>Natural selection<\/strong>: A consistent bias in survival and fertility, leading to the overrepresentation of certain features in future generations and an improved fit between an average member of the population and the environment.<\/p>\n<p class=\"import-Normal\"><strong>Niche construction<\/strong>: The active engagement by which species transform their surroundings in favorable ways, rather than just passively inhabiting them.<\/p>\n<p class=\"import-Normal\"><strong>Phenotype<\/strong>: Observable manifestation of a genetic constitution, expressed in a particular set of circumstances. The suite of traits of an organism.<\/p>\n<p class=\"import-Normal\"><strong>Phrenology<\/strong>: The 19th-century anatomical study of bumps on the head as an indication of personality and mental abilities.<\/p>\n<p class=\"import-Normal\"><strong>Plasticity<\/strong>: The tendency of a growing organism to react developmentally to its particular conditions of life.<\/p>\n<p class=\"import-Normal\"><strong>Punctuated equilibria<\/strong>: The idea that species are stable through time and are formed very rapidly relative to their duration. (The opposite theory, that species are unstable and constantly changing through time, is called phyletic gradualism.)<\/p>\n<p class=\"import-Normal\"><strong>Scientific racism<\/strong>: The use of pseudoscientific evidence to support or legitimize racial hierarchy and inequality.<\/p>\n<p class=\"import-Normal\"><strong>Sexual selection<\/strong>: Natural selection arising through preference by one sex for certain characteristics in individuals of the other sex.<\/p>\n<p class=\"import-Normal\"><strong>Species selection<\/strong>: A postulated evolutionary process in which selection acts on an entire species population, rather than individuals.<\/p>\n<h2 class=\"import-Normal\">About the Authors<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image2-4-1.jpg\" alt=\"A bearded man wearing glasses smiles at the camera. \" width=\"202\" height=\"218\" \/><\/p>\n<h3 class=\"import-Normal\"><strong>Jonathan Marks, Ph.D.<\/strong><\/h3>\n<p class=\"import-Normal\">University of North Carolina at Charlotte, <a class=\"rId41\" href=\"mailto:jmarks@uncc.edu\">jmarks@uncc.edu<\/a><\/p>\n<p class=\"import-Normal\">Jonathan Marks is Professor of Anthropology at the University of North Carolina at Charlotte. He has published many books and articles on broad aspects of biological anthropology. In 2006 he was elected a Fellow of the American Association for the Advancement of Science. In 2012 he was awarded the First Citizen\u2019s Bank Scholar\u2019s Medal from UNC Charlotte. In recent years he has been a Visiting Research Fellow at the ESRC Genomics Forum in Edinburgh, a Visiting Research Fellow at the Max Planck Institute for the History of Science in Berlin, and a Templeton Fellow at the Institute for Advanced Study at Notre Dame. His work has received the W. W. Howells Book Prize and the General Anthropology Division Prize for Exemplary Cross-Field Scholarship from the American Anthropological Association as well as the J. I. Staley Prize from the School for Advanced Research. Two of his books are titled <em>What It Means to Be 98% Chimpanzee<\/em> and <em>Why I Am Not a Scientist<\/em>, but actually he is about 98 percent scientist and not a chimpanzee.<\/p>\n<p class=\"import-Normal\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image10-3.jpg\" alt=\"A bearded man wearing a fedora hat looks off in the distance. \" width=\"232\" height=\"232\" \/><\/p>\n<h3 class=\"import-Normal\"><strong>Adam P. Johnson, M.A.<\/strong><\/h3>\n<p class=\"import-Normal\">University of North Carolina at Charlotte\/University of Texas at San Antonio, <a class=\"rId43\" href=\"mailto:ajohn344@uncc.edu\">ajohn344@uncc.edu<\/a><\/p>\n<p class=\"import-Normal\">Adam Johnson is a doctoral candidate at the University of Texas at San Antonio and part-time lecturer at the University of North Carolina at Charlotte. He earned his M.A. in anthropology at UNC-Charlotte in 2017 and will complete his Ph.D. in anthropology at UTSA by 2024. His interests include human-animal relations, science studies, primate behavior, ecology, and the history of anthropology. His recent research project analyzes the social, historical, political, and evolutionary dimensions that shape human-javelina encounters. His goal is to understand how humans and animals find ways to get along in a precarious world.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration <strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\">Ackermann, Rebecca Rogers, Alex Mackay, and Michael L. Arnold. 2016. \u201cThe Hybrid Origin of \u2018Modern\u2019 Humans.\u201d <em>Evolutionary Biology<\/em> 43 (1): 1\u201311.<\/p>\n<p class=\"import-Normal\">Bateson, Patrick, and Peter Gluckman. 2011. <em>Plasticity, Robustness, Development and Evolution<\/em>. New York: Cambridge University Press.<\/p>\n<p class=\"import-Normal\">Cosans, Christopher E. 2009. <em>Owen's Ape and Darwin's Bulldog: Beyond Darwinism and Creationism<\/em>. Bloomington, IN: Indiana University Press.<\/p>\n<p class=\"import-Normal\">Desmond, Adrian, and James Moore. 2009. <em>Darwin's Sacred Cause: How a Hatred of Slavery Shaped Darwin's Views on Human Evolution<\/em>. New York: Houghton Mifflin Harcourt.<\/p>\n<p class=\"import-Normal\">Dobzhansky, Theodosius, Francisco J. Ayala, G. Ledyard Stebbins, and James W. Valentine. 1977. <em>Evolution<\/em>. San Francisco: W.H. Freeman and Company.<\/p>\n<p class=\"import-Normal\">Fuentes, Agust\u00edn. 2017. <em>The Creative Spark: How Imagination Made Humans Exceptional<\/em>. New York: Dutton.<\/p>\n<p class=\"import-Normal\">Gould, Stephen J. 2003.<em> The Structure of Evolutionary Theory<\/em>. Cambridge, MA: Harvard University Press.<\/p>\n<p class=\"import-Normal\">Haraway, Donna J. 1989. <em>Primate Visions: Gender, Race, and Nature in the World of Modern Science<\/em>. New York: Routledge.<\/p>\n<p class=\"import-Normal\">Huxley, Thomas. 1863. <em>Evidence as to Man's Place in Nature<\/em>. London: Williams &amp; Norgate.<\/p>\n<p class=\"import-Normal\">Jablonka, Eva, and Marion J. Lamb. 2005. <em>Evolution in Four Dimensions: Genetic, Epigenetic, Behavioral, and Symbolic Variation in the History of Life<\/em>. Cambridge, MA: The MIT Press.<\/p>\n<p class=\"import-Normal\">Kuklick, Henrika, ed. 2008. <em>A New History of Anthropology<\/em>. New York: Blackwell.<\/p>\n<p class=\"import-Normal\">Laland, Kevin N., Tobias Uller, Marcus W. Feldman, Kim Sterelny, Gerd B. Muller, Armin Moczek, Eva Jablonka, and John Odling-Smee. 2015. \u201cThe Extended Evolutionary Synthesis: Its Structure, Assumptions and Predictions.\u201d <em>Proceedings of the Royal Society, Series B<\/em> 282 (1813): 20151019.<\/p>\n<p class=\"import-Normal\">Lamarck, Jean Baptiste. 1809. <em>Philosophie Zoologique<\/em>. Paris: Dentu.<\/p>\n<p class=\"import-Normal\">Landau, Misia. 1991. <em>Narratives of Human Evolution<\/em>. New Haven: Yale University Press.<\/p>\n<p class=\"import-Normal\">Lee, Sang-Hee. 2017. <em>Close Encounters with Humankind: A Paleoanthropologist Investigates Our Evolving Species<\/em>. New York: W. W. Norton.<\/p>\n<p class=\"import-Normal\">Livingstone, David N. 2008. <em>Adam's Ancestors: Race, Religion, and the Politics of Human Origins<\/em>. Baltimore: Johns Hopkins University Press.<\/p>\n<p class=\"import-Normal\">Marks, Jonathan. 2015. <em>Tales of the Ex-Apes: How We Think about Human Evolution<\/em>. Berkeley, CA: University of California Press.<\/p>\n<p class=\"import-Normal\">Pigliucci, Massimo. 2009. \u201cThe Year in Evolutionary Biology 2009: An Extended Synthesis for Evolutionary Biology.\u201d <em>Annals of the New York Academy of Sciences<\/em> 1168: 218\u2013228.<\/p>\n<p class=\"import-Normal\">Simpson, George Gaylord. 1949. <em>The Meaning of Evolution: A Study of the History of Life and of Its Significance for Man<\/em>. New Haven: Yale University Press.<\/p>\n<p class=\"import-Normal\">Sommer, Marianne. 2016.<em> History Within: The Science, Culture, and Politics of Bones, Organisms, and Molecules<\/em>. Chicago: University of Chicago Press.<\/p>\n<p class=\"import-Normal\">Stoczkowski, Wiktor. 2002. <em>Explaining Human Origins: Myth, Imagination and Conjecture<\/em>. New York: Cambridge University Press.<\/p>\n<p class=\"import-Normal\">Tattersall, Ian, and Rob DeSalle. 2019. <em>The Accidental Homo sapiens: Genetics, Behavior, and Free Will<\/em>. New York: Pegasus.<\/p>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\">Barton, Robert A. 1996. \"Neocortex Size and Behavioural Ecology in Primates.\" <em>Proceedings of the Royal Society of London. Series B: Biological Sciences<\/em> 263 (1367): 173\u2013177.<\/p>\n<p class=\"import-Normal\">Bodmer, Walter, and Robin McKie. 1997. <em>The Book of Man: The Hman Genome Project and the Quest to Discover our Genetic Heritage.<\/em> Oxford University Press.<\/p>\n<p class=\"import-Normal\">Darwin, Charles. 1859.<em> On the Origin of Species by Means of Natural Selection, or, the Preservation of Favoured Races in the Struggle for Life<\/em>. London: J. Murray.<\/p>\n<p class=\"import-Normal\">Darwin, Charles. 1871. <em>The Descent of Man, and Selection in Relation to Sex.<\/em> London: J. Murray.<\/p>\n<p class=\"import-Normal\">Dawkins, Richard. 1976. <em>The Selfish Gene. <\/em>Oxford University Press.<\/p>\n<p class=\"import-Normal\">Deacon, T. W. 1998. <em>The Symbolic Species: The Co-evolution of Language and the Brain<\/em>. W. W. Norton &amp; Company.<\/p>\n<p class=\"import-Normal\">Eldredge, N., and S. J. Gould. 1972. \"Punctuated Equilibria: An Alternative to Phyletic Gradualism.\" In <em>Models in Paleobiology<\/em>, edited by T. J. Schopf, 82\u2013115. San Francisco: W. H. Freeman.<\/p>\n<p class=\"import-Normal\">Gould, Stephen J. 2003.<em> The Structure of Evolutionary Theory<\/em>. Cambridge, MA: Harvard University Press.<\/p>\n<p class=\"import-Normal\">Gould, Stephen J. 1996. <em>Mismeasure of Man<\/em>. New York: WW Norton &amp; Company.<\/p>\n<p class=\"import-Normal\">Gould, Stephen Jay, and Richard C. Lewontin. 1979. \"The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme.\" <em>Proceedings of the Royal Society of London. Series B: Biological Sciences<\/em> 205 (1151): 581\u2013598.<\/p>\n<p class=\"import-Normal\">Haeckel, Ernst. 1868. <em>Nat\u00fcrliche Sch\u00f6pfungsgeschichte<\/em>. Berlin: Reimer.<\/p>\n<p class=\"import-Normal\">Huxley, Thomas Henry. 1863. <em>Evidence as to Man\u2019s Place in Nature. <\/em>London: Williams and Norgate.<\/p>\n<p class=\"import-Normal\">Kaufman, Thomas C., Mark A. Seeger, and Gary Olsen. 1990. \"Molecular and Genetic Organization of the Antennapedia Gene Complex of <em>Drosophila melanogaster<\/em>.\" <em>Advances in Genetics<\/em> 27: 309\u2013362.<\/p>\n<p class=\"import-Normal\">Kellogg, Vernon. 1917. <em>Headquarters Nights<\/em>. Boston: The Atlantic Monthly Press.<\/p>\n<p class=\"import-Normal\">Kevles, Daniel J., and Leroy Hood. 1993. <em>The Code of Codes: Scientific and Social Issues in the Human Genome Project<\/em>. Cambridge, MA: Harvard University Press.<\/p>\n<p class=\"import-Normal\">Lewontin, Richard, Steven Rose, and Leon Kamin. 2017. <em>Not in Our Genes\u202f: Biology, Ideology, and Human Nature<\/em>, 2nd ed. Chicago: Haymarket Books.<\/p>\n<p class=\"import-Normal\">Lloyd, Elisabeth A., and Stephen J. Gould. 1993. \"Species Selection on Variability.\" <em>Proceedings of the National Academy of Sciences<\/em> 90 (2): 595\u2013599.<\/p>\n<p class=\"import-Normal\">Marks, Jonathan. 2015. \u201cThe Biological Myth of Human Evolution.\u201d In <em>Biologising the Social Sciences: Challenging Darwinian and Neuroscience Explanations<\/em>, edited by David Canter and David A. Turner, 59\u201378. London: Routledge.<\/p>\n<p class=\"import-Normal\">Monypenny, William Flavelle, and George Earle Buckle. 1929. <em>The Life of Benjamin Disraeli, Earl of Beaconsfield, Volume II: 1860\u20131881<\/em>. London: John Murray.<\/p>\n<p class=\"import-Normal\">Potts, Rick. 1998. \u201cVariability Selection in Hominid Evolution.\u201d <em>Evolutionary Anthropology <\/em><em>7<\/em><em>:<\/em> 81\u201396.<\/p>\n<p class=\"import-Normal\">Punnett, R. C. 1905. <em>Mendelism<\/em>. Cambridge: Macmillan and Bowes.<\/p>\n<p class=\"import-Normal\">Shapiro, Robert. 1991. <em>The Human Blueprint: The Race to Unlock the Secrets of Our Genetic Script.<\/em> New York: St. Martin\u2019s Press.<\/p>\n<p class=\"import-Normal\">Shultz, Susanne, Emma Nelson, and Robin Dunbar. 2012. \"Hominin Cognitive Evolution: Identifying Patterns and Processes in the Fossil and Archaeological Record.\" <em>Philosophical Transactions of the Royal Society B: Biological Sciences<\/em> 367 (1599): 2130\u20132140.<\/p>\n<p class=\"import-Normal\">Spencer, Herbert. 1864. <em>Principles of Biology.<\/em> London: Williams and Norgate.<\/p>\n<p class=\"import-Normal\">Watson, James D. 1990. \"The Human Genome Project: Past, Present, and Future.\" <em>Science<\/em> 248 (4951): 44\u201349.<\/p>\n<p class=\"import-Normal\">Yengo, L., Vedantam, S., Marouli, E., Sidorenko, J., Bartell, E., Sakaue, S., Graff, M., Eliasen, A.U., Jiang, Y., Raghavan, S. and Miao, J., 2022. A saturated map of common genetic variants associated with human height. <em>Nature<\/em>, <em>610 <\/em>(7933): 704-712.<\/p>\n<p class=\"import-Normal\">Zeder, Melinda A. 2018. \"Why Evolutionary Biology Needs Anthropology: Evaluating Core Assumptions of the Extended Evolutionary Synthesis.\" <em>Evolutionary Anthropology: Issues, News, and Reviews<\/em> 27 (6): 267\u2013284.<\/p>\n<\/div>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_2709\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_2709\"><div tabindex=\"-1\"><\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1802\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1802\"><div tabindex=\"-1\"><\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1760\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1760\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\">Jonathan Marks, Ph.D., University of North Carolina at Charlotte<\/p>\n<p class=\"import-Normal\">Adam P. Johnson, M.A., University of North Carolina at Charlotte\/University of Texas at San Antonio<\/p>\n<p class=\"import-Normal\"><em>This chapter is an adaptation of \"<\/em><a class=\"rId9\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__\/\"><em>Chapter 2: Evolution<\/em><\/a><em>\u201d by Jonathan Marks. In <\/em><a class=\"rId10\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, first edition<\/em><\/a><em>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under <\/em><a class=\"rId11\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\"><em>CC BY-NC 4.0<\/em><\/a><em>. <\/em><\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\"><span style=\"color: #ffffff\">Learning Objectives<\/span><\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li>Explain the relationship among genes, bodies, and organismal change.<\/li>\n<li>Discuss the shortcomings of simplistic understandings of genetics.<\/li>\n<li>Describe what is meant by the \"biopolitics of heredity.\"<\/li>\n<li>Discuss issues caused by misuse of ideas about adaptations and natural selection.<\/li>\n<li>Examine and correct myths about evolution.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p class=\"import-Normal\">The Human Genome Project, an international initiative launched in 1990, sought to identify the entire genetic makeup of our species. For many scientists, it meant trying to understand the genetic underpinnings of what made humans uniquely human. James Watson, a codiscoverer of the helical shape of DNA, wrote that \u201cwhen finally interpreted, the genetic messages encoded within our DNA molecules will provide the ultimate answers to the chemical underpinnings of human existence\u201d (Watson 1990, 248). The underlying message is that what makes humans unique can be found in our <strong>genes<\/strong>. The Human Genome Project hoped to find the core of who we are and where we come from.<\/p>\n<p class=\"import-Normal\">Despite its lofty goal, the Human Genome Project\u2014even after publishing the entire human genome in January 2022\u2014could not fully account for the many factors that contribute to what it is to be human. Richard Lewontin, Steven Rose, and Leon Kamin (2017) argue that genetic determinism of the sort assumed by the Human Genome Project neglects other essential dimensions that contribute to the development and evolution of human bodies, not to mention the role that culture plays. They use an apt metaphor of a cake to illustrate the incompleteness of reductive models. Consider the flavor of a cake and think of the ingredients listed in the recipe. The recipe includes ingredients such as flour, sugar, shortening, vanilla extract, eggs, and milk. Does raw flour taste like cake? Does sugar, vanilla extract, or any of the other ingredients taste like cake? They do not, and knowing the individual flavors of each ingredient does not tell us much about what cake tastes like. Even mixing all of the ingredients in the correct proportions does not get us cake. Instead, external factors such as baking at the right temperature, for the right amount of time, and even the particularities of our evolved sense of taste and smell are all necessary components of experiencing the cake.<\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ff00ff\">Lewontin, Rose, and Kamin (2017) argue that the same is true for humans and other organisms.<\/span><\/p>\n<p class=\"import-Normal\">Knowing everything about cake ingredients does not allow us to fully know cake. Equally so, knowing everything about the genes found in our DNA does not allow us to fully know humans. Different, interacting levels are implicated in the development and evolution of all organisms, including humans. Genes, the structure of chromosomes, developmental processes, epigenetic tags, environmental factors, and still-other components all play key roles such that genetically reductive models of human development and evolution are woefully inadequate.<\/p>\n<p class=\"import-Normal\">The complex interactions across many levels\u2014genetic, developmental, and environmental\u2014explain why we still do not know how our one-dimensional DNA nucleotide sequence results in a four-dimensional organism. This was the unfulfilled promise of the inception of the Human Genome Project in the 1980s and 1990s: the project produced the complete DNA sequence of a human cell in the hopes that it would reveal how human bodies are built and how to cure them when they are built poorly. Yet, that information has remained elusive. Presumably, the knowledge of how organisms are produced from DNA sequences will one day permit us to reconcile the discrepancies between patterns in anatomical evolution and molecular evolution.<\/p>\n<p class=\"import-Normal\">In this chapter, we will consider multilevel evolution and explore evolution as a complex interaction between genetic and epigenetic factors as well as the environments in which organisms live. Next, we will examine the biopolitical nature of human evolution. We will then investigate problems that arise from attributing all traits to an adaptive function. Finally, we will address common misconceptions about evolution. The goal of this chapter is to provide you with the necessary toolkit for understanding the molecular, anatomical, and political dimensions of evolution.<\/p>\n<h2 class=\"import-Normal\">Evolution Happens at Multiple Levels<\/h2>\n<p class=\"import-Normal\">Following Richard Dawkins\u2019s publication of <em>The Selfish Gene <\/em>in 1976, the scientific imagination was captured by the potential of genomics to reveal how genes are copied by Darwinian selection. Dawkins argues that the genes in individuals that contribute to greater reproductive success are the units of selection. His conception of evolution at the molecular level undercuts the complex interactions between organisms and their environments, which are not expressed genomically but are nevertheless key drivers in evolution.<\/p>\n<p class=\"import-Normal\">By the 1980s, the acknowledgment among most biologists that even though genes construct bodies, genes and bodies evolve at different rates and with distinct patterns. This realization led to a renewed focus on how bodies change. The Evolutionary Synthesis of the 1930s\u20131970s had reduced organisms to their <strong>genotypes<\/strong> and species to their <strong>gene pools<\/strong>, which provided valuable insights about the processes of biological change, but it was only a first approximation. Animals are in fact reactive and adaptable beings, not passive and inert genotypes. Species are clusters of socially interacting and reproductively compatible organisms.<\/p>\n<figure style=\"width: 291px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/06\/image8-5.png\" alt=\"An asteroid hits the ocean. Pterodactyls fly among clouds in the foreground.\" width=\"291\" height=\"233\" \/><figcaption class=\"wp-caption-text\">Figure 17.1: A painting by Donald E. Davis representing the Chicxulub asteroid impact off the Yucatan Peninsula that contributed to the mass extinction that included the dinosaurs about 65 million years ago. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Chicxulub_impact_-_artist_impression.jpg\">Chicxulub impact - artist impression<\/a> by Donald E. Davis, <a href=\"https:\/\/www.nasa.gov\/\">NASA<\/a>, is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Once we accept that evolutionary change is fundamentally genetic change, we can ask: How do bodies function and evolve? How do groups of animals come to see one another as potential mates or competitors for mates, as opposed to just other creatures in the environment? Are there evolutionary processes that are not explicable by population genetics? These questions\u2014which lead us beyond reductive assumptions\u2014were raised in the 1980s by Stephen Jay Gould, the leading evolutionary biologist of the late 20th century (see: Gould 2003; 1996).<\/p>\n<p class=\"import-Normal\">Gould spearheaded a movement to identify and examine higher-order processes and features of evolution that were not adequately explained by population genetics. For example, <strong>extinction<\/strong>, which was such a problem for biologists of the 1600s, could now be seen as playing a more complex role in the history of life than population genetics had been able to model. Gould recognized that there are two kinds of extinctions, each with different consequences: background extinctions and mass extinctions. Background extinctions are those that reflect the balance of nature, because in a competitive Darwinian world, some things go extinct and other things take their place. Ecologically, your species may be adapted to its niche, but if another species comes along that\u2019s better adapted to the same niche, eventually your species will go extinct. It sucks, but it is the way of all life: you come into existence, you endure, and you pass out of existence. But mass extinctions are quite different. They reflect not so much the balance of nature as the wholesale disruption of nature: many species from many different lineages dying off at roughly the same time\u2014presumably as the result of some kind of rare ecological disaster. The situation may not be survival of the fittest as much as survival of the luckiest. The result, then, would be an ecological scramble among the survivors. Having made it through the worst, the survivors could now simply divide up the new ecosystem amongst themselves, since their competitors were gone. Something like this may well have happened about 65 million years ago, when a huge asteroid hit the Yucatan Peninsula, which mammals survived but dinosaurs did not (Figure 17.1). Something like this may be happening now, due to human expansion and environmental degradation. Note, though, that there is only a limited descriptive role here for population genetics: the phenomena we are describing are about organisms and species in ecosystems.<\/p>\n<p class=\"import-Normal\">Another question involved the disconnect between properties of <em>species<\/em> and the properties of <em>gene pools<\/em>. For example, there are upwards of 15 species of gibbons but only two species of chimpanzees. Why? There are upwards of 20 species of guenons but fewer than ten of baboons. Why? Are there genes for that? It seems unlikely. Gould suggested that species, as units of nature, might have properties that are not reducible to the genes in their cells. For example, rates of speciation and extinction might be properties of their ecologies and histories rather than their genes. Thus, relationships between environmental contexts and variability within a species result in degrees of resistance to extinction and affect the frequency and rates at which clades diversify (Lloyd and Gould 1993). Consistent biases of speciation rates might well produce patterns of macroevolutionary diversity that are difficult to explain genetically and better understood ecologically. Gould called such biases in speciation rates <strong>species selection<\/strong>\u2014a higher-order process that invokes competition between species, in addition to the classic Darwinian competition between individuals.<\/p>\n<p class=\"import-Normal\">One of Gould\u2019s most important studies involved the very nature of species. In the classical view, a species is continually adapting to its environment until it changes so much that it is a different species than it was at the beginning of this sentence (Eldredge and Gould 1972). That implies that the species is a fundamentally unstable entity through time, continuously changing to fit in. But suppose, argued Gould along with paleontologist Niles Eldredge, a species is more stable through time and only really adapts during periods of ecological instability and change. Then we might expect to find in the fossil record long equilibrium periods\u2014a few million years or so\u2014in which species don\u2019t seem to change much, punctuated by relatively brief periods in which they change a bit and then stabilize again as new species. They called this idea <strong>punctuated equilibria<\/strong>. The idea helps to explain certain features of the fossil record, notably the existence of small anatomical \u201cgaps\u201d between closely related fossil forms (Figure 17.2). Its significance lies in the fact that although it incorporates genetics, punctuated equilibria is not really a theory of genetics but one of types bodies in deep time.<\/p>\n<p class=\"import-Normal\">Punctuated equilibria is seen across taxa, with long periods in the fossil record representing little phenotypic change. These periods of stability are disrupted by shorter periods of rapid <strong>adaptation<\/strong>, the process through which populations of organisms become suited to living in their environments. Phenotypic changes are often coupled with drastic climatic or ecological changes that affect the milieu in which organisms live. For example, throughout much of hominin evolutionary history, brain size was closely associated with body size and thus remained mostly stable. However, changes occurred in average hominin brain size at around 100 thousand years ago, 1 million years ago, and 1.8 million years ago. Several hypotheses have been put forth to explain these changes, including unpredictability in climate and environment (Potts 1998), social development (Barton 1996), and the evolution of language (Deacon 1998). Evidence from the fossil record, paleoclimate models, and comparative anatomy suggests that the changes observed in hominin lineage result from biocultural processes\u2014that is, the coalescence of environmental and cultural factors that selected for larger brains (Marks 2015; Shultz, Nelson, and Dunbar 2012).<\/p>\n<figure style=\"width: 461px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image5-8.png\" alt=\"Two graphs contrast phyletic gradualism and punctuated equilibria.\" width=\"461\" height=\"222\" \/><figcaption class=\"wp-caption-text\">Figure 17.2: Different ways of conceptualizing the evolutionary relationship between an earlier and a later species. With phyletic gradualism, species are envisioned transforming continually in a direct line over time. With punctuated equilibria species branch off at particular points over time.\u00a0 Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__\/\">Phyletic gradualism vs. punctuated equilibria (Figure 2.12)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">In response to the call for a theory of the evolution of form, the field of <strong>evo-devo<\/strong>\u2014the intersection of evolutionary and developmental biology\u2014arose. The central focus here is on how changes in form and shape arise. An embryo matures by the stimulation of certain cells to divide, forming growth fields. The interactions and relationships among these growth fields generate the structures of the body. The <strong>hox genes<\/strong> that regulate these growth fields turn out to be highly conserved across the animal kingdom. This is because they repeatedly turn on and off the most basic genes guiding the animal\u2019s development, and thus any changes to them would be catastrophic. Indeed, these genes were first identified by manipulating them in fruit flies, such that one could produce a bizarre mutant fruit fly that grew a pair of legs where its antennae were supposed to be (Kaufman, Seeger, and Olsen 1990).<\/p>\n<p class=\"import-Normal\">Certain genetic changes can alter the fates of cells and the body parts, while other genetic changes can simply affect the rates at which neighboring groups of cells grow and divide, thus producing physical bumps or dents in the developing body. The result of altering the relationships among these fields of cellular proliferation in the growing embryo is <strong>allometry<\/strong>, or the differential growth of body parts. As an animal gets larger\u2014either over the course of its life or over the course of macroevolution\u2014it often has to change shape in order to live at a different size. Many important physiological functions depend on properties of geometric area: the strength of a bone, for example, is proportional to its cross-sectional area. But area is a two-dimensional quality, while growing takes place in three dimensions\u2014as an increase in mass or volume. As an animal expands, its bones necessarily weaken, because volume expands faster than area does. Consequently a bigger animal has more stress on its bones than a smaller animal does and must evolve bones even thicker than they would be by simply scaling the animal up proportionally. In other words, if you expand a mouse to the size of an elephant, it will nevertheless still have much thinner bones than the elephant does. But those giant mouse bones will unfortunately not be adequate to the task. Thus, a giant mouse would have to change aspects of its form to maintain function at a larger size (see Figure 17.3).<\/p>\n<p class=\"import-Normal\"><img class=\"aligncenter\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image6-6.png\" alt=\"Side-view of a mouse skeleton.\" width=\"515\" height=\"252\" \/><\/p>\n<figure style=\"width: 453px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image7-9.png\" alt=\"Side-view of an elephant skeleton.\" width=\"453\" height=\"326\" \/><figcaption class=\"wp-caption-text\">Figure 17.3: Mouse (top) and elephant (bottom) skeletons. Notice the elephant\u2019s bones are more robust when the two animals are the same size. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__\/\">Mouse and elephant skeletons (Figure 2.13)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Physiologically, we would like to know how the body \u201cknows\u201d when to turn on and off the genes that regulate growth to produce a normal animal. Evolutionarily, we would like to know how the body \u201clearns\u201d to alter the genetic on\/off switch (or the genetic \u201cslow down\/speed up\u201d switch) to produce an animal that looks different. Moreover, since organisms differ from one another, we would like to know how the developing body distinguishes a range of normal variation from abnormal variation. And, finally, how does abnormal variation eventually become normal in a descendant species?<\/p>\n<p class=\"import-Normal\">Taking up these questions, Gould invoked the work of a British geneticist named Conrad H. Waddington, who thought about genetics in less reductive ways than his colleagues. Rather than isolate specific DNA sites to analyze their function, Waddington instead studied the inheritance of an organism\u2019s reactivity\u2014its ability to adapt to the circumstances of its life. In a famous experiment, he grew fruit fly eggs in an atmosphere containing ether. Most died, but a few survived somehow by developing a weird physical feature: a second thorax with a second pair of wings. Waddington bred these flies and soon developed a stable line of flies who would reliably develop a second thorax when grown in ether. Then he began to lower the concentration of ether, while continuing to selectively breed the flies that developed the strange appearance. Eventually he had a line of flies that would stably develop the \u201cbithorax\u201d <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1238\"><strong>phenotype<\/strong><\/a>\u2013the suite of traits of an organism\u2013even when there was no ether; it had become the \u201cnew normal.\u201d The flies had genetically assimilated the bithorax condition.<\/p>\n<p class=\"import-Normal\">Waddington was thus able to mimic the <strong>inheritance of acquired characteristics<\/strong>: what had been a trait stimulated by ether a few generations ago was now a normal part of the development of the descendants. Waddington recognized that while he had performed a selection experiment on genetic variants, he had not selected for particular traits. Rather, he helped produce the physiological tendency to develop particular traits when appropriately stimulated. He called that tendency <strong>plasticity<\/strong> and its converse, the tendency to stay the same even under weird environmental circumstances, <strong>canalization.<\/strong> Waddington had initially selected for plasticity, the tendency to develop the bithorax phenotype under weird conditions, and then, later, for canalization, the developmental normalization of that weird physical trait. Although Waddington had high stature in the community of geneticists, evolutionary biologists of the 1950s and 1960s regarded him with suspicion because he was not working within the standard mindset of reductionism, which saw evolution as the spread of genetic variants that coded for favorable traits. Both Waddington and Gould resisted contemporary intellectual paradigms that favored reductive accounts of evolutionary processes. They conceived of evolution as an emergent process in which many external factors (e.g. climate, environment, predation) and internal factors (e.g., genotypes, plasticity, canalization) coalesce to produce the evolutionary trends that we observe in the fossil record and our genome.<\/p>\n<p class=\"import-Normal\">While Gould and Waddington both looked beyond the genome to understand evolution, the Human Genome Project\u2014an international project with the goal of identifying each base pair in the human genome in the 1990s\u2014generated a great deal of public interest in analyzing the human DNA sequence from the standpoint of medical genetics. Some of the rhetoric aimed to sell the public on investing a lot of money and resources in sequencing the human genome in order to show the genetic basis of heritable traits, cure genetic diseases, and learn what it means ultimately to be biologically human. However, the Human Genome Project was not actually able to answer those questions through the use of genetics alone, and thus a broader, more holistic account was required.<\/p>\n<p class=\"import-Normal\">This holistic account came from decades of research in human biology and anthropology, which understood the human body as highly adaptable, dynamic, and emergent. For example, in the early 20th century, anthropologist Franz Boas measured the skulls of immigrants to the U.S., revealing that environmental, not merely genetic, factors affected skull shape. The growing human body adjusts itself to the conditions of life, such as diet, sunshine, high altitude, hard labor, population density, how babies are carried\u2014any and all of which can have subtle but consistent effects upon its development. There can thus be no normal human form, only a context-specific range of human forms.<\/p>\n<p class=\"import-Normal\">However, what the human biologists called human adaptability, evolutionary biologists called developmental plasticity, and evidence quickly began to mount for its cause being <strong>epigenetic <\/strong>modifications to DNA. Epigenetic modifications are changes to how genes are used by the body (as opposed to changes in the DNA sequences; see Chapter 3). Scientific interest shifted from the focus of the Human Genome Project to the ways that bodies are made by evolutionary-developmental processes, including epigenetics. What is meant by \u201cepigenetic modification\u201d? Evolution is about how descendants diverge from their ancestors. Inheritance from parent to offspring is still critical to this process, which occurs through genetic recombination: the pairing of homologous chromosomes and sharing of genetic material during meiosis (see Chapter 3). However, in the 21st century, the link between evolution and inheritance has broadened with a clearer understanding of how environmental and developmental factors shape bodies and the expression of genes, including epigenetic inheritance patterns. While offspring inherit their genes through random assortment during meiosis, environmental factors also shape how genes are used. When these epigenetic modifications occur in germ cells, they can be passed onto offspring. In these cases, there is no change in the DNA sequence but rather in how genes are used by the body due to DNA methylation and the structure of chromosomes due to histone acetylation (see Chapter 3).<\/p>\n<p class=\"import-Normal\">In addition, we now recognize that evolution is affected by two other forms of intergenerational transmission and inheritance (in addition to genetics and epigenetics). These forms include behavioral variation and culture. That is, behavioral information can be transmitted horizontally (intragenerationally), permitting more rapid ways for organisms to adjust to the environment. And, then there is the fourth mode of transmission: the cultural or symbolic mode. <span style=\"background-color: #ffff00\">Humans are the only species<\/span> that horizontally transmits an arbitrary set of rules to govern communication, social interaction, and thought. This shared information is symbolic and has resulted in what we recognize as \u201cculture\u201d: locally emergent worlds of names, words, pictures, classifications, revered pasts, possible futures, spirits, dead ancestors, unborn descendants, in-laws, politeness, taboo, justice, beauty, and story, all accompanied by practices and a material world of tools.<\/p>\n<p class=\"import-Normal\">Consequently our contemporary ideas about evolution see the evolutionary processes as hierarchically organized and not restricted to the differential transmission of DNA sequences into the next generation. While that is indeed a significant part of evolution, the organism and species are nevertheless crucial to understanding how those DNA sequences get transmitted. Further, the transmission of epigenetic, behavioral, and symbolic information play a complex role in perpetuating our genes, bodies, and species. In the case of human evolution, one can readily see that symbolic information and cultural adaptation are far more central to our lives and our survival today than DNA and genetic adaptation. It is thus misleading to think of humans passively occupying an environmental niche. Rather, humans are actively engaged in constructing our own niches, as well as adapting to them and using them to adapt. The complex interplay between a species and its active engagement in creating its own ecology is known as <strong>niche construction<\/strong>. If we understand <strong>natural selection<\/strong>\u2013the process by which populations adapt to their specific environments\u2013as the effects that environmental context has on the reproductive success of organisms, then niche construction is the process through which organisms shape their own selective pressures.<\/p>\n<h2 class=\"import-Normal\">The Biopolitics of Heredity<\/h2>\n<p class=\"import-Normal\">\u201cScience isn\u2019t political\u201d is a sentiment that you have likely heard before. Science is supposed to be about facts and objectivity. It exists, or at least ought to, outside of petty human concerns. However, the sorts of questions we ask as scientists, the problems we choose to study, the categories and concepts we use, who gets to do science, and whose work gets cited are all shaped by culture. Doing science is a political act. This fact is markedly true for human evolution. While it is easier to create intellectual distance between us and fruit flies and viruses, there is no distance when we are studying ourselves. The hardest lesson to learn about human evolution is that it is intensely political. Indeed, to see it from the opposite side, as it were, the history of creationism\u2014the belief that the universe was divinely created around 6,000 years ago\u2014is essentially a history of legal decisions. For instance, in <em>Tennessee v. John T. Scopes<\/em> (1925), a schoolteacher was prosecuted for violating a law in Tennessee that prohibited the teaching of human evolution in public schools, where teachers were required by law to teach creationism.<\/p>\n<p class=\"import-Normal\">More recently, legal decisions aimed at legislating science education have shaped how students are exposed to evolutionary theory. For instance, <em>McLean v. Arkansas<\/em> (1982) dispatched \u201cscientific creationism\u201d by arguing that the imposition of balanced teaching of evolution and creationism in science classes violates the Establishment Clause, separating church and state. Additionally, <em>Kitzmiller v. Dover (Pennsylvania) Area School District<\/em> (2005) dispatched the teaching of \u201cintelligent design\u201d in public school classrooms as it was deemed to not be science. In some cases, people see unbiblical things in evolution, although most Christian theologians are easily able to reconcile science to the Bible. In other cases, people see immoral things in evolution, although there is morality and immorality everywhere. And some people see evolution as an aspect of alt-religion, usurping the authority of science in schools to teach the rejection of the Christian faith, which would be unconstitutional due to the protected separation of church and state.<\/p>\n<p class=\"import-Normal\">Clearly, the position that politics has nothing to do with science is untenable. But is the politics in evolution an aberration or is it somehow embedded in science? In the early 20th century, scientists commonly promoted the view that science and politics were separate: science was seen as a pure activity, only rarely corrupted by politics. And yet as early as World War I, the politics of nationalism made a hero of the German chemist Fritz Haber for inventing poison gas. And during World War II, both German doctors and American physicists, recruited to the war effort, helped to end many civilian lives. Therefore, we can think of the apolitical scientist as a self-serving myth that functions to absolve scientists of responsibility for their politics. The history of science shows how every generation of scientists has used evolutionary theory to rationalize political and moral positions. In the very first generation of evolutionary science, Darwin\u2019s <em>Origin of Species<\/em> (1859) is today far more readable than his <em>Descent of Man<\/em> (1871). The reason is that Darwin consciously purged <em>The Origin of Species<\/em> of any discussion of people. And when he finally got around to talking about people, in <em>The Descent of Man<\/em>, he simply imbued them with the quaint Victorian prejudices of his age, and the result makes you cringe every few pages. There is plenty of politics in there\u2014sexism, racism, and colonialism\u2014because <em>you cannot talk about people apolitically<\/em>.<\/p>\n<p class=\"import-Normal\">One immediate faddish deduction from Darwinism, popularized by Herbert Spencer (1864) as \u201csurvival of the fittest,\u201d held that unfettered competition led to advancement in nature and to human history. Since the poor were purported losers in that struggle, anything that made their lives easier would go against the natural order. This position later came to be known ironically as \u201cSocial Darwinism.\u201d Spencer was challenged by fellow Darwinian Thomas Huxley (1863), who agreed that struggle was the law of the jungle but observed that we don\u2019t live in jungles anymore. The obligation to make lives better for others is a moral, not a natural, fact. We simultaneously inhabit a natural universe of descent from apes and a moral universe of injustice and inequality, and science is not well served by ignoring the latter.<\/p>\n<p class=\"import-Normal\">Concurrently, the German biologist Ernst Haeckel\u2019s 1868 popularization of Darwinism was translated into English a few years later as <em>The History of Creation<\/em>. As we saw earlier, Haeckel was determined to convince his readers that they were descended from apes, even in the absence of fossil evidence attesting to it. When he made non-Europeans into the missing links that connected his readers to the apes, and depicted them as ugly caricatures, he knew precisely what he was doing. Indeed, even when the degrading racial drawings were deleted from the English translation of his book, the text nevertheless made his arguments quite clear. And a generation later, when the Americans had not yet entered the Great War in 1916, a biologist named Vernon Kellogg visited the German High Command as a neutral observer and found that the officers knew a lot about evolutionary biology, which they had gotten from Haeckel and which rationalized their military aggressions. Kellogg went home and wrote a bestseller about it, called <em>Headquarters Nights<\/em> (1917). World War I would have been fought with or without evolutionary theory, but as a source of scientific authority, evolution\u2014even if a perversion of the Darwinian theory\u2014had very quickly attained global geopolitical relevance.<\/p>\n<p class=\"import-Normal\">Oftentimes, politics in evolutionary science is subtle, due to the pervasive belief in the advancement of science. We recognize the biases of our academic ancestors and modify our scientific stories accordingly. But we can never be free of our own cultural biases, which are invisible to us, as much as our predecessors\u2019 biases were invisible to them. In some cases, the most important cultural issues resurface in different guises each generation, like scientific racism. <strong>Scientific racism<\/strong> is the recruitment of science for the evil political ends of racism, and it has proved remarkably impervious to evolution. Before Darwin, there was creationist scientific racism, and after Darwin, there was evolutionist scientific racism. And there is still scientific racism today, self-justified by recourse to evolution, which means that scientists have to be politically astute and sensitive to the uses of their work to counter these social tendencies.<\/p>\n<p class=\"import-Normal\">Consider this: Are you just your ancestry, or can you transcend it? If that sounds like a weird question, it was actually quite important to a turn-of-the-20th-century European society in which an old hereditary aristocracy was under increasing threat from a rising middle class. And that is why the very first English textbook of Mendelian genetics concluded with the thought that \u201cpermanent progress is a question of breeding rather than of pedagogics; a matter of gametes, not of training \u2026 the creature is not made but born\u201d (Punnett 1905, 60). <em>Translation: Not only do we now know a bit about how heredity works, but it\u2019s also the most important thing about you. Trust me, I\u2019m a scientist.<\/em><\/p>\n<p class=\"import-Normal\">Yet evolution is about how descendants come to differ from ancestors. Do we really know that your heredity, your DNA, your ancestry, is the most important thing about you? That you were born, not made? After all, we do know that you could be born into slavery or as a peasant, and come from a long line of enslaved people or peasants, and yet not have slavery or peasantry be the most important thing about you. Whatever your ancestors were may unfortunately constrain what you can become, but as a moral precept, it should not. But just as science is not purely \u201cfacts and objectivity,\u201d ancestry is not a strictly biological concept. Human ancestry is biopolitics, not biology.<\/p>\n<p class=\"import-Normal\">Evolution is fundamentally a theory about ancestry, and yet ancestors are, in the broad anthropological sense, sacred: ancestors are often more meaningful symbolically than biologically. Just a few years after <em>The Origin of Species <\/em>(Darwin 1859), the British politician and writer Benjamin Disraeli declared himself to be on the side of the angels, not the apes, and to \u201crepudiate with indignation and abhorrence those new-fangled theories\u201d (Monypenny, Flavelle, and Buckle 1920, 105). He turned his back on an ape ancestry and looked to the angel; yet, he did so as a prominent Jew-turned-Anglican, who had personally transcended his humble roots and risen to the pinnacle of the Empire. Ancestry was certainly important, and Disraeli was famously proud of his, but it was also certainly not the most important thing, not the primary determinant of his place in the world. Indeed, quite the opposite: Disraeli\u2019s life was built on the transcendence of many centuries of Jewish poverty and oppression in Europe. Humble ancestry was there to be superseded and nobility was there to be earned; Disraeli would later become the Earl of Beaconsfield. Clearly, \u201care you just your ancestry\u201d is not a value-neutral question, and \u201cthe creature is not made, but born\u201d is not a value-neutral answer.<\/p>\n<p class=\"import-Normal\">Ancestry being the most important thing about a person became a popular idea twice in 20th century science. First, at the beginning of the century, when the <strong>eugenics<\/strong> movement in America called attention to \u201cfeeble-minded stocks,\u201d which usually referred to the poor or to immigrants (see Figure 17.4; and see Chapter 2). This movement culminated in Congress restricting the immigration of \u201cfeeble-minded races\u201d (said to include Jews and Italians) in 1924, and the Supreme Court declaring it acceptable for states to sterilize their \u201cfeeble-minded\u201d citizens involuntarily in 1927. After the Nazis picked up and embellished these ideas during World War II, Americans moved swiftly away from them in some contexts (e.g., for most people of European descent) while still strictly adhering in other contexts (e.g., Japanese internment camps and immigration restrictions).<\/p>\n<figure style=\"width: 374px\" class=\"wp-caption alignright\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image4-6.png\" alt=\"Historic photo. People sit in front of a structure with a \u201cEugenic and Health Exhibit&quot; banner.\" width=\"374\" height=\"262\" \/><figcaption class=\"wp-caption-text\">Figure 17.4: Eugenic and Health Exhibit, Fitter Families exhibit, and examination building, Kansas State Free Fair. Credit: <a href=\"https:\/\/www.dnalc.org\/view\/16328-Gallery-14-Eugenics-Exhibit-at-the-Kansas-State-Free-Fair-1920.html\">Gallery 14: Eugenics Exhibit at the Kansas State Free Fair, 1920 ID (ID 16328)<\/a> by <a href=\"https:\/\/www.dnalc.org\/\">Cold Spring Harbor<\/a> (Courtesy American Philosophical Society) is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/3.0\/us\/\">CC BY-NC-ND 3.0 License.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\">Ancestry again became paramount in the drumming up of public support for the Human Genome Project in the 1990s. Public support for sequencing the human genome was encouraged by a popular science campaign that featured books titled <em>The Book of Man <\/em>(Bodmer and McKie 1997), <em>The Human Blueprint <\/em>(Shapiro 1991), and <em>The Code of Codes<\/em> (Kevles and Hood 1993). These books generally promised cures for genetic diseases and a deeper understanding of the human condition. We can certainly identify progress in molecular genetics over the last couple of decades since the human genome was sequenced, but that progress has notably not been accompanied by cures for genetic diseases, nor by deeper understandings of the human condition.<\/p>\n<p class=\"import-Normal\">Even at the most detailed and refined levels of genetic analysis, we still don\u2019t have much of an understanding of the actual basis by which things seem to \u201crun in families.\u201d While the genetic basis of simple, if tragic, genetic diseases have become well-known\u2014such as sickle-cell anemia, cystic fibrosis, and Tay-Sachs\u2019 Disease\u2014we still haven\u2019t found the ostensible genetic basis for traits that are thought to have a strong genetic component. For example, a recent genetic summary found over 12,000 genetic sites that contributed to height yet still explained only about 40-50 percent of the variation in height among European ancestry but no more than 10-20 percent of variation of other ancestries, which we know strongly runs in families (Yengo et al. 2022).<\/p>\n<p class=\"import-Normal\">Partly in reaction to the reductionistic hype of the Human Genome Project, the study of epigenetics has become the subject of great interest. One famous natural experiment involves a Nazi-imposed famine in Holland over the winter of 1944\u20131945. Children born during and shortly after the famine experienced a higher incidence of certain health problems as adults, many decades later. Apparently, certain genes had been down-regulated early in development and remained that way throughout the course of life. Indeed, this modified regulation of the genes in response to the severe environmental conditions may have been passed on to their children.<\/p>\n<p class=\"import-Normal\">Obviously one\u2019s particular genetic constitution may play an important role in one\u2019s life trajectory. But overvaluing that role may have important social and political consequences. In the first place, genotypes are rendered meaningful in a cultural universe. Thus, if you live in a strongly patriarchal society and are born without a Y chromosome (since human males are chromosomally XY and females XX), your genotype will indeed have a strong effect upon your life course. So even though the variation is natural, the consequences are political. The mediating factors are the cultural ideas about how people of different sexes ought to be treated, and the role of the state in permitting certain people to develop and thrive. More broadly, there are implications for public education if variation in intelligence is genetic. There are implications for the legal system if criminality is genetic. There are implications for the justice system if sexual preference, or sexual identity, is genetic. There are implications for the development of sports talent if that is genetic. And yet, even for the human traits that are more straightforward to measure and known to be strongly heritable, the DNA base sequence variation seems to explain little.<\/p>\n<p class=\"import-Normal\">Genetic determinism or <strong>hereditarianism<\/strong> is the idea that \u201cthe creature is made, not born\u201d\u2014or, in a more recent formulation by James Watson, that \u201cour fate is in our genes.\u201d One of the major implications drawn from genetic determinism is that the feature in question must inevitably express itself; therefore, we can\u2019t do anything about it. Therefore, we might as well not fund the social programs designed to ameliorate economic inequality and improve people\u2019s lives, because their courses are fated genetically. And therefore, they don\u2019t deserve better lives.<\/p>\n<p class=\"import-Normal\">All of the \u201ctherefores\u201d in the preceding paragraph are open to debate. What is important is that the argument relies on a very narrow understanding of the role of genetics in human life, and it misdirects the causes of inequality from cultural to natural processes. By contrast, instead of focusing on genes and imagining them to place an invisible limit upon social progress, we can study the ways in which your DNA sequence does <em>not<\/em> limit your capability for self-improvement or fix your place in a social hierarchy. In general, two such avenues exist. First, we can examine the ways in which the human body responds and reacts to environmental variation: human adaptability and plasticity. This line of research began with the anthropometric studies of immigrants by Franz Boas in the early 20th century and has now expanded to incorporate the epigenetic inheritance of modified human DNA. And second, we can consider how human lives are shaped by social histories\u2014especially the structural inequalities within the societies in which they grow up.<\/p>\n<p class=\"import-Normal\">Although it arises and is refuted every generation, the radical hereditarian position (genetic determinism) perennially claims to speak for both science and evolution. It does not. It is the voice of a radical fringe\u2014perhaps naive, perhaps evil. It is not the authentic voice of science or of evolution. Indeed, keeping Charles Darwin\u2019s name unsullied by protecting it from association with bad science often seems like a full-time job. Culture and epigenetics are very much a part of the human condition, and their roles are significant parts of the complete story of human evolution.<\/p>\n<p><span style=\"background-color: #00ffff\"><span style=\"text-decoration: underline\">(Sterilization of Indigenous women in Canada)<\/span> (https:\/\/www.thecanadianencyclopedia.ca\/en\/article\/sterilization-of-indigenous-women-in-canada)\u00a0<\/span><\/p>\n<h2 class=\"import-Normal\">Adaptationism and the Panglossian Paradigm<\/h2>\n<p class=\"import-Normal\">The story of human evolution, and the evolution of all life for that matter, is never settled because evolution is ongoing. Additionally, because the conditions that shape evolutionary trajectories are not predetermined, evolution itself is emergent. Even during periods of ecological stability, when fewer macroevolutionary changes occur, populations of organisms continue to experience change. When ecological stability is disrupted, populations must adapt to the changes. Darwin explained in naturalistic terms how animals adapt to their environments: traits that contribute to an organism's ability to survive and reproduce in specific environments will become more common. The most \u201cfit\u201d\u2014those organisms best suited to the <em>current<\/em> environmental conditions in which they live\u2014have survived over eons of the history of life on earth to cocreate ecosystems full of animals and plants. Our own bodies are full of evident adaptations: eyes for seeing, ears for hearing, feet for walking on, and so forth.<\/p>\n<p class=\"import-Normal\">But what about hands? Feet are adapted to be primarily weight-bearing structures (rather than grasping structures, as in the apes) and that is what we primarily use them for. But we use our hands in many ways: for fine-scale manipulation, greeting, pointing, stimulating a sexual partner, writing, throwing, and cooking, among other uses. So which of these uses express what hands are \u201cfor,\u201d when all of them express what hands do?<\/p>\n<p class=\"import-Normal\">Gould and Lewontin (1979) illustrate the problem with assuming that the function of a trait defines its evolutionary cause. Consider the case of Dr. Pangloss\u2014the protagonistic of Voltaire\u2019s <em>Candide<\/em>\u2014who believed that we lived in the best of all possible worlds. Gould and Lewontin use his pronouncement that \u201cnoses were made for spectacles and so we have spectacles\u201d to demonstrate the problem with assuming any trait has evolved for a specific purpose. Identifying a function of a trait does not necessitate that the function is the ultimate cause of the trait. Individual traits are not under selection pressures in isolation; in fact, an entire organism must be able to survive and reproduce in their environment. When natural selection results in adaptations, changes that occur in some traits can have cascading effects throughout the phenotype and features that are not under selection pressure can also change.<\/p>\n<figure style=\"width: 279px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image3-5.png\" alt=\"Human hand is smaller with smaller fingers and smoother skin compared to a chimpanzee hand.\" width=\"279\" height=\"264\" \/><figcaption class=\"wp-caption-text\">Figure 17.5: Drawings of a human hand (left) and a chimpanzee hand (right). Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__\/\">Human and chimpanzee hand (Figure 2.16)<\/a> by Mary Nelson original to <a href=\"https:\/\/explorations.americananthro.org\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">There is an important lesson in recognizing that what things do in the present is not a good guide to understanding why they came to exist. Gunpowder was invented for entertainment\u2014only later was it adopted for killing people. The Internet was invented to decentralize computers in case of a nuclear attack\u2014and only later adopted for social media. Apes have short thumbs and use their hands in locomotion; our ancestors stopped using their hands in locomotion by about six million years ago and had fairly modern-looking hands by about two million years ago. We can speculate that a combination of selection for abstract thought and dexterity led to evolution of the human hand, with its capability for toolmaking that exceeds what apes can do (see Figure 17.5). But let\u2019s face it\u2014how many tools have you made today?<\/p>\n<p class=\"import-Normal\">Consequently, we are obliged to see the human foot as having a purpose to which it is adapted and the human hand as having multiple purposes, most of which are different from what it originally evolved for. Paleontologists Gould and Elisabeth Vrba suggested that an original use be regarded as an adaptation and any additional uses be called \u201c<strong>exaptations.<\/strong>\u201d Thus, we would consider the human hand to be an adaptation for toolmaking and an exaptation for writing. So how do we know whether any particular feature is an adaptation, like the walking foot, rather than an exaptation, like the writing hand? Or more broadly, how can we reason rigorously from what a feature does to what it evolved for?<\/p>\n<p class=\"import-Normal\">The answer to the question \u201cwhat did this feature evolve for?\u201d creates an origin myth. This origin myth contains three assumptions: (1) features can be isolated as evolutionary units; (2) there is a specific reason for the existence of any particular feature; and (3) there is a clear and simplistic explanation for why the feature evolved.<\/p>\n<figure style=\"width: 378px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image9-8.png\" alt=\"Head with images and human qualities drawn on it. Journal title printed at the bottom.\" width=\"378\" height=\"437\" \/><figcaption class=\"wp-caption-text\">Figure 17.6: According to the early 19th century science of phrenology, units of personality could be mapped onto units in the head, as shown on this cover of The Phrenology Journal. Credit: <a href=\"https:\/\/wellcomecollection.org\/works\/b6skynug\">Phrenology; Chart<\/a> [slide number 5278, photo number: L0000992, original print from Dr. E. Clark, The Phrenological Journal (Know Thyself)] by <a href=\"https:\/\/wellcomecollection.org\/\">Wellcome Collection<\/a>, is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/legalcode\">CC BY 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">The first assumption was appreciated a century ago as the \u201cunit-character problem.\u201d Are the units by which the body grows and evolves the same as units we name? This is clearly not the case: we have genes and we have noses, and we have genes that affect noses, but we don\u2019t have \u201cnose genes.\u201d What is the relationship between the evolving elements that we see, identify, and name, and the elements that biologically exist and evolve? It is hard to know, but we can use the history of science as a guide to see how that fallacy has been used by earlier generations. Back in the 19th century, the early anatomists argued that since the brain contained the mind, they could map different mental states (acquisitiveness, punctuality, sensitivity) onto parts of the brain. Someone who was very introspective, say, would have an enlarged introspection part of the brain, a cranial bulge to represent the hyperactivity of this mental state. The anatomical science was known as <strong>phrenology<\/strong>, and it was predicated on the false assumption that units of thought or personality or behavior could be mapped to distinct parts of the brain and physically observed (see Figure17.6). This is the fallacy of reification, imagining that something named is something real.<\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Long alt text: Side view of human head. At the top are the words \u201cKnow Thyself.\u201d On the upper head are small illustrations and word qualities such as \u201cfriendship,\u201d \u201cself-esteem,\u201d and \u201csecretiveness.\u201d On the lower part of the man\u2019s man\u2019s face are the words <em>The Phrenological Journal and Science of Health, A First Class Monthly<\/em>. The caption at the bottom reads: \u201cSpecially devoted to the \u2018.\u2019 Contains PHRENOLOGY and PHYSIOGNOMY, with all the SIGNS OF CHARACTER, and how to read them; ETHNOLOGY, or the Natural History of Man in all his relations.\u201d (All emphases in original.)<\/span><\/p>\n<figure style=\"width: 295px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image1-8-1.png\" alt=\"A black-and-white drawing of a chimpanzee head and face.\" width=\"295\" height=\"236\" \/><figcaption class=\"wp-caption-text\">Figure 17.7: Chimpanzees have big ears. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Chimpanzee_head_sketch.png\">Chimpanzee head sketch<\/a> by <a href=\"https:\/\/de.wikipedia.org\/wiki\/Benutzer:Roger_Zenner\">Roger Zenner<\/a>, original by Brehms Tierleben (1887), is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">The second assumption, that everything has a reason, has long been recognized as a core belief of religion. Our desire to impose order and simplicity on the workings of the universe, however, does not constrain it to obey simple and orderly causes. Magic, witchcraft, spirits, and divine agency are all powerful explanations for why things happen. Consequently, it is probably not a good idea to lump natural selection in with those. Sometimes things do happen for a reason, of course, but other times things happen as byproducts of other things, or for very complicated and entangled reasons, or for no reason at all. What phenomena have reasons and thereby merit explanation? Chimpanzees have very large testicles, and we think we know why: their promiscuous sexual behavior triggers intense competition for high sperm count. But chimpanzees also have very large ears, but much less scientific attention has been paid to this trait (see Figure 17.7). Why not? Why should there be a reason for chimp testicles but not for chimp ears? What determines the kinds of features that we try to explain, as opposed to the ones that we do not? Again, the assumption that any specific feature has a reason is metaphysical; that is to say, it may be true in any particular case, but to assume it in all cases is gratuitous.<\/p>\n<p class=\"import-Normal\">And third, the possibility of knowing what the reason for any particular feature is, assuming that it has one, is a challenge for evolutionary epistemology (the theory of how we know things). Consider the big adaptations of our lineage: bipedalism and language. Nobody doubts that they are good, and they evolved by natural selection, and we know how they work. But why did they evolve? If talking and walking are simply better than not talking and not walking, then why did they evolve in just a single branch of the ape lineage in the primate family tree? We don\u2019t know what bipedalism evolved for, although there are plenty of speculations: walking long distances, running long distances, cooling the head, seeing over tall grass, carrying babies, carrying food, wading, threatening, counting calories, sexual display, and so on. Neither do we know what language evolved for, although there are speculations: coordinating hunting, gossiping, manipulating others. But it is also possible that bipedality is simply the way that a small arboreal ape travels on the ground, if it isn\u2019t in the treetops. Or that language is simply the way that a primate with small canine teeth and certain mental propensities comes to communicate. If that were true, then there might be no reason for bipedality or language: having the unique suite of preconditions and a fortuitous set of circumstances simply set them in motion, and natural selection elaborated and explored their potentials. It is possible that walking and talking simply solved problems that no other lineage had ever solved; but even if so, the fact remains that the rest of the species in the history of life have done pretty well without having solved them.<\/p>\n<p class=\"import-Normal\">It is certainly very optimistic to think that all three assumptions (that organisms can be meaningfully atomized, that everything has a reason, and that we can know the reason) would be simultaneously in effect. Indeed, just as there are many ways of adapting (genetically, epigenetically, behaviorally, culturally), there are also many ways of being nonadaptive, which would imply that there is no reason at all for the feature in question.<\/p>\n<p class=\"import-Normal\">First, there is the element of randomness of population histories. There are more cases of sickle-cell anemia among sub-Saharan Africans than other peoples, and there is a reason for it: carriers of sickle-cell anemia have a resistance to malaria, which is more frequent in parts of Africa (as discussed in Chapters 4 and 14). But there are more cases of a blood disease called variegated porphyria, a rare genetic metabolic disorder, in the Afrikaners of South Africa (descendants of mostly Dutch settlers in the 17th century) than in other peoples, and there is no reason for it. Yet we know the cause: One of the founding Dutch colonial settlers had the <strong>allele<\/strong>\u2013a variant of a gene\u2013and everyone in South Africa with it today is her descendant. But that is not a reason\u2014that is simply an accident of history.<\/p>\n<p class=\"import-Normal\">Second, there is the potential mismatch between the past and the present. The value of a particular feature in the past may be changed as the environmental circumstances change. Our species is diurnal, and our ancestors were diurnal. But beginning around a few hundred thousand years ago, our ancestors could build fires, which extended the light period, which was subsequently further amplified by lamps and candles. And over the course of the 20th century, electrical power has made it possible for people to stay up very late when it is dark\u2014working, partying, worrying\u2014to a greater extent than any other closely related species. In other words, we evolved to be diurnal, yet we are now far more nocturnal than any of our recent ancestors or close relatives. Are we adapting to nocturnality? If so, why? Does it even make any sense to speak of the human occupation of a nocturnal ape niche, despite the fact that we empirically seem to be doing just that? And if so, does it make sense to ask what the reason for it is?<\/p>\n<p class=\"import-Normal\">Third, there is a genetic phenomenon known as a selective sweep, or the hitchhiker effect. Imagine three genes\u2014A, B, and C\u2014located very closely together on a chromosome. They each have several variants, or alleles, in the population. Now, for whatever reason, it becomes beneficial to have one of the B alleles, say B4; this B4 allele is now under strong positive selection. Obviously, we will expect future generations to be characterized by mostly B4. But what was B4 attached to? Because whatever A and C alleles were adjacent to it will also be quickly spread, simply by virtue of the selection for B4. Even if the A and C alleles are not very good, they will spread because of the good B4 allele between them. Eventually the linkage groups will break up because of genetic crossing-over in future generations. But in the meantime, some random version of genes A and C are proliferating in the species simply because they are joined to superior allele B4. And clearly, the A and C alleles are there because of selection\u2014but not because of selection <em>for<\/em> them!<\/p>\n<p class=\"import-Normal\">Fourth, some features are simply consequences of other properties rather than adaptations to external conditions. We already noted the phenomenon of allometric growth, in which some physical features have to outgrow others to maintain function at an increased size. Can we ask the reason for the massive brow ridges of <em>Homo erectus<\/em>, or are brow ridges simply what you get when you have a conjunction of thick skull bones, a large face, and a sloping forehead\u2014and, thus, again would have a cause but no reason?<\/p>\n<p class=\"import-Normal\">Fifth, some features may be underutilized and on the way out. What is the reason for our two outer toes? They aren\u2019t propulsive, they don\u2019t do anything, and sometimes they\u2019re just in the way. Obviously they are there because we are descended from ancestors with five digits on their hands and feet. Is it possible that a million years from now, we will just have our three largest toes, just as the ancestors of the horse lost their digits in favor of a single hoof per limb? Or will our outer toes find another use, such as stabilizing the landings in our personal jet-packs? For the time being, we can just recognize vestigiality as another nonadaptive explanation for the presence of a given feature.<\/p>\n<p class=\"import-Normal\">Finally, Darwin himself recognized that many obvious features do not help an animal survive. Some things may instead help an animal breed. The peacock\u2019s tail feathers do not help it eat, but they do help it mate. There is competition, but only against half of the species. Darwin called this <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1084\">sexual selection<\/a><\/strong>. Its result is not a fit to the environment but, rather, a fit to the opposite sex. In some species, that is literally the case, as the male and female genitalia have specific ways of anatomically fitting together. The specific form is less important than the specific match, so inquiring about the reason for a particular form of the reproductive anatomy may be misleading. The specific form may be effectively random, as long as it fits the opposite sex and is different from the anatomies of other species. Nor is sexual selection the only form of selection that can affect the body differently from natural selection. Competition might also take place between biological units other than organisms\u2014perhaps genes, perhaps cells, or populations, or species. The spread of cultural things, such as head-binding or cheap refined fructose or forced labor, can have significant effects upon bodies, which are also not adaptations produced by natural selection. They are often adaptive physiological responses to stresses but not the products of natural selection.<\/p>\n<p class=\"import-Normal\">With so many paths available by which a physical feature might have organically arisen without having been the object of natural selection, it is unwise to assume that any individual trait is an adaptation. And that generalization applies to the best-known, best-studied, and most materially based evolutionary adaptations of our lineage. But our cultural behaviors are also highly adaptive, so what about our most familiar social behaviors? Patriarchy, hierarchy, warfare\u2014are these adaptations? Do they have reasons? Are they good for something?<\/p>\n<p class=\"import-Normal\">This is where some sloppy thinking has been troublesome. What would it mean to say that patriarchy evolved by natural selection in the human species? If, on the one hand, it means that the human mind evolved by natural selection to be able to create and survive in many different kinds of social and political regimes, of which patriarchy is one, then biological anthropologists will readily agree. If, on the other hand, it means that patriarchy evolved by natural selection, that implies that patriarchy is genetically determined (since natural selection is a genetic process) and out-reproduced the alleles for other, more egalitarian, social forms. This in turn would imply that patriarchy is an adaptation and therefore of some beneficial value in the past and has become an ingrained part of human nature today. This would be bad news, say, if you harbored ambitions of dismantling it. Dismantling patriarchy in that case would be to go against nature, a futile gesture. In other words, this latter interpretation would be a naturalistic manifesto for a conservative political platform: don\u2019t try to dismantle the patriarchy, because it is within us, the product of evolution\u2014suck it up and live with it.<\/p>\n<p class=\"import-Normal\">Here, evolution is being used as a political instrument for transforming the human genome into an imaginary glass ceiling against equality. There is thus a convergence between the pseudo-biology of crude <strong>adaptationism <\/strong>(the idea that everything is the product of natural selection) and the pseudo-biology of hereditarianism. Naturalizing inequality is not the business of evolutionary theory, and it represents a difficult moral position for a scientist to adopt, as well as a poor scientific position.<\/p>\n<h3><strong>Evolution of the Anthropocene (to be reviewed)<\/strong><\/h3>\n<p>As humans have caused the emergence of the Anthropocene, it is important to inform scholars about the effect of our social and cultural evolution on the rest of the world. Richard Robbins\u2019 <em>Global Problems and Culture of Capitalism<\/em> explains how the modern culture of consumption has been extremely successful at accommodating populations of people far larger than previously possible. Robbins claims that the globalization attributed to capitalism has allowed the world to make full use of its environmental resources, providing necessities and innovative technologies to humans all over the world (Robbins &amp; Dowty, 2019). In other words, capitalism is an anthropocentric cultural system that highly benefits humans and facilitates our survival with little regard to the development and survival of other forms of life. It would be of high relevance to introduce the idea that our cultural evolution and capacity to modify the environment to meet our needs have established new environmental conditions in which the human species' survival and reproduction rate expand at the detriment of ecosystems and endangerment of other primates and non-human species.<\/p>\n<p>According to a 2019 UN report, following the 16th century, the world has entered a period of extreme environmental destruction that is generating ecological modifications and has led to the extinction of at least 680 vertebrate species and over 9 percent of the domesticated mammals used for food and agriculture (United Nations, 2019). Human lifestyles are causing changes that\u2014if not taken into consideration\u2014could lead to our extinction as a species. The recognition that our evolutionary behavioural development is causing environmental destruction may be the first step for our species to take accountability for the damage that it is causing to others and prevent further damage.<\/p>\n<p>&nbsp;<\/p>\n<h2 class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Concluding Thoughts<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Now that you have finished reading this chapter, you are equipped to understand the historical and political dimensions of evolution. Evolution is an ongoing process of change and diversification. Evolutionary theory is a tool that we use to understand this process. The development of evolutionary theory is shaped both by scientific innovation and political engagement. Since Darwin first articulated natural selection as an observable mechanism by which species adapt to their environments, our understanding of evolution has grown. Initially, scientists focused on the adaptive aspects of evolution. However, with the emergence of genetics, our understanding of heredity and the level at which evolution acts has changed. Genetics led to a focus on the molecular dimensions of evolution. For some, this focus resulted in reductive accounts of evolution. Further developments in our understanding of evolution shifted our view to epigenetic processes and how organisms shape their own evolutionary pressures (e.g., niche construction).<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Evolutionary theory will continue to develop in the future as we invent new technologies, describe new dimensions of biology, and experience cultural changes. Current innovations in evolutionary theory are asking us to consider evolutionary forces beyond natural selection and genetics to include the ways organisms shape their environments (niche construction), inheritances beyond genetics (inclusive inheritance), constraints on evolutionary change (developmental bias), and the ability of bodies to change in response to external factors (plasticity). The future of evolutionary theory looks bright as we continue to explore these and other dimensions. Biological anthropology is well-positioned to be a lively part of this conversation, as it extends standard evolutionary theory by considering the role of culture, social learning, and human intentionality in shaping the evolutionary trajectories of humans (Zeder 2018). Remember, at root, human evolutionary theory consists of two propositions: (1) the human species is descended from other similar species and (2) natural selection has been the primary agent of biological adaptation. Pretty much everything else is subject to some degree of contestation.<\/span><\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Review Questions<\/h2>\n<ul>\n<li class=\"import-Normal\">How is the study of your ancestors biopolitical, not just biological? Does that make it less scientific or differently scientific?<\/li>\n<li class=\"import-Normal\">What was gained by reducing organisms to genotypes and species to gene pools? What is gained by reintroducing bodies and species into evolutionary studies?<\/li>\n<li class=\"import-Normal\">How do genetic or molecular studies complement anatomical studies of evolution?<\/li>\n<li class=\"import-Normal\">How are you reducible to your ancestry? If you could meet your ancestors from the year 1700 (and you would have well over a thousand of them!), would their lives be meaningfully similar to yours? Would you even be able to communicate with them?<\/li>\n<li class=\"import-Normal\">The molecular biologist Fran\u00e7ois Jacob argued that evolution is more like a tinkerer than an engineer. In what ways do we seem like precisely engineered machinery, and in what ways do we seem like jerry-rigged or improvised contraptions?<\/li>\n<li class=\"import-Normal\">How might biological anthropology contribute to future developments in evolutionary theory?<\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\">Key Terms<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\"><strong>Adaptation<\/strong>: A fit between the organism and environment.<\/p>\n<p class=\"import-Normal\"><strong>Adaptationism<\/strong>: The idea that everything is the product of natural selection.<\/p>\n<p class=\"import-Normal\"><strong>Allele<\/strong>: A genetic variant.<\/p>\n<p class=\"import-Normal\"><strong>Allometry<\/strong>: The differential growth of body parts.<\/p>\n<p class=\"import-Normal\"><strong>Canalization<\/strong>: The tendency of a growing organism to be buffered toward normal development.<\/p>\n<p class=\"import-Normal\"><strong>Epigenetics<\/strong>: The study of how genetically identical cells and organisms (with the same DNA base sequence) can nevertheless differ in stably inherited ways.<\/p>\n<p class=\"import-Normal\"><strong>Eugenics<\/strong>: An idea that was popular in the 1920s that society should be improved by breeding \u201cbetter\u201d kinds of people.<\/p>\n<p class=\"import-Normal\"><strong>Evo-devo<\/strong>: The study of the origin of form; a contraction of \u201cevolutionary developmental biology.\u201d<\/p>\n<p class=\"import-Normal\"><strong>Exaptation<\/strong>: An additional beneficial use for a biological feature.<\/p>\n<p class=\"import-Normal\"><strong>Extinction<\/strong>: The loss of a species from the face of the earth.<\/p>\n<p class=\"import-Normal\"><strong>Gene<\/strong>: A stretch of DNA with an identifiable function (sometimes broadened to include any DNA with recognizable structural features as well).<\/p>\n<p class=\"import-Normal\"><strong>Gene pool<\/strong>: Hypothetical summation of the entire genetic composition of population or species.<\/p>\n<p class=\"import-Normal\"><strong>Genotype<\/strong>: Genetic constitution of an individual organism.<\/p>\n<p class=\"import-Normal\"><strong>Hereditarianism<\/strong>: The idea that genes or ancestry is the most crucial or salient element in a human life. Generally associated with an argument for natural inequality on pseudo-genetic grounds.<\/p>\n<p class=\"import-Normal\"><strong>Hox genes<\/strong>: A group of related genes that control for the body plan of an embryo along the head-tail axis.<\/p>\n<p class=\"import-Normal\"><strong>Inheritance of acquired characteristics<\/strong>: The idea that you pass on the features that developed during your lifetime, not just your genes; also known as Lamarckian inheritance.<\/p>\n<p class=\"import-Normal\"><strong>Natural selection<\/strong>: A consistent bias in survival and fertility, leading to the overrepresentation of certain features in future generations and an improved fit between an average member of the population and the environment.<\/p>\n<p class=\"import-Normal\"><strong>Niche construction<\/strong>: The active engagement by which species transform their surroundings in favorable ways, rather than just passively inhabiting them.<\/p>\n<p class=\"import-Normal\"><strong>Phenotype<\/strong>: Observable manifestation of a genetic constitution, expressed in a particular set of circumstances. The suite of traits of an organism.<\/p>\n<p class=\"import-Normal\"><strong>Phrenology<\/strong>: The 19th-century anatomical study of bumps on the head as an indication of personality and mental abilities.<\/p>\n<p class=\"import-Normal\"><strong>Plasticity<\/strong>: The tendency of a growing organism to react developmentally to its particular conditions of life.<\/p>\n<p class=\"import-Normal\"><strong>Punctuated equilibria<\/strong>: The idea that species are stable through time and are formed very rapidly relative to their duration. (The opposite theory, that species are unstable and constantly changing through time, is called phyletic gradualism.)<\/p>\n<p class=\"import-Normal\"><strong>Scientific racism<\/strong>: The use of pseudoscientific evidence to support or legitimize racial hierarchy and inequality.<\/p>\n<p class=\"import-Normal\"><strong>Sexual selection<\/strong>: Natural selection arising through preference by one sex for certain characteristics in individuals of the other sex.<\/p>\n<p class=\"import-Normal\"><strong>Species selection<\/strong>: A postulated evolutionary process in which selection acts on an entire species population, rather than individuals.<\/p>\n<h2 class=\"import-Normal\">About the Authors<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image2-4-1.jpg\" alt=\"A bearded man wearing glasses smiles at the camera. \" width=\"202\" height=\"218\" \/><\/p>\n<h3 class=\"import-Normal\"><strong>Jonathan Marks, Ph.D.<\/strong><\/h3>\n<p class=\"import-Normal\">University of North Carolina at Charlotte, <a class=\"rId41\" href=\"mailto:jmarks@uncc.edu\">jmarks@uncc.edu<\/a><\/p>\n<p class=\"import-Normal\">Jonathan Marks is Professor of Anthropology at the University of North Carolina at Charlotte. He has published many books and articles on broad aspects of biological anthropology. In 2006 he was elected a Fellow of the American Association for the Advancement of Science. In 2012 he was awarded the First Citizen\u2019s Bank Scholar\u2019s Medal from UNC Charlotte. In recent years he has been a Visiting Research Fellow at the ESRC Genomics Forum in Edinburgh, a Visiting Research Fellow at the Max Planck Institute for the History of Science in Berlin, and a Templeton Fellow at the Institute for Advanced Study at Notre Dame. His work has received the W. W. Howells Book Prize and the General Anthropology Division Prize for Exemplary Cross-Field Scholarship from the American Anthropological Association as well as the J. I. Staley Prize from the School for Advanced Research. Two of his books are titled <em>What It Means to Be 98% Chimpanzee<\/em> and <em>Why I Am Not a Scientist<\/em>, but actually he is about 98 percent scientist and not a chimpanzee.<\/p>\n<p class=\"import-Normal\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image10-3.jpg\" alt=\"A bearded man wearing a fedora hat looks off in the distance. \" width=\"232\" height=\"232\" \/><\/p>\n<h3 class=\"import-Normal\"><strong>Adam P. Johnson, M.A.<\/strong><\/h3>\n<p class=\"import-Normal\">University of North Carolina at Charlotte\/University of Texas at San Antonio, <a class=\"rId43\" href=\"mailto:ajohn344@uncc.edu\">ajohn344@uncc.edu<\/a><\/p>\n<p class=\"import-Normal\">Adam Johnson is a doctoral candidate at the University of Texas at San Antonio and part-time lecturer at the University of North Carolina at Charlotte. He earned his M.A. in anthropology at UNC-Charlotte in 2017 and will complete his Ph.D. in anthropology at UTSA by 2024. His interests include human-animal relations, science studies, primate behavior, ecology, and the history of anthropology. His recent research project analyzes the social, historical, political, and evolutionary dimensions that shape human-javelina encounters. His goal is to understand how humans and animals find ways to get along in a precarious world.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration <strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\">Ackermann, Rebecca Rogers, Alex Mackay, and Michael L. Arnold. 2016. \u201cThe Hybrid Origin of \u2018Modern\u2019 Humans.\u201d <em>Evolutionary Biology<\/em> 43 (1): 1\u201311.<\/p>\n<p class=\"import-Normal\">Bateson, Patrick, and Peter Gluckman. 2011. <em>Plasticity, Robustness, Development and Evolution<\/em>. New York: Cambridge University Press.<\/p>\n<p class=\"import-Normal\">Cosans, Christopher E. 2009. <em>Owen's Ape and Darwin's Bulldog: Beyond Darwinism and Creationism<\/em>. Bloomington, IN: Indiana University Press.<\/p>\n<p class=\"import-Normal\">Desmond, Adrian, and James Moore. 2009. <em>Darwin's Sacred Cause: How a Hatred of Slavery Shaped Darwin's Views on Human Evolution<\/em>. New York: Houghton Mifflin Harcourt.<\/p>\n<p class=\"import-Normal\">Dobzhansky, Theodosius, Francisco J. Ayala, G. Ledyard Stebbins, and James W. Valentine. 1977. <em>Evolution<\/em>. San Francisco: W.H. Freeman and Company.<\/p>\n<p class=\"import-Normal\">Fuentes, Agust\u00edn. 2017. <em>The Creative Spark: How Imagination Made Humans Exceptional<\/em>. New York: Dutton.<\/p>\n<p class=\"import-Normal\">Gould, Stephen J. 2003.<em> The Structure of Evolutionary Theory<\/em>. Cambridge, MA: Harvard University Press.<\/p>\n<p class=\"import-Normal\">Haraway, Donna J. 1989. <em>Primate Visions: Gender, Race, and Nature in the World of Modern Science<\/em>. New York: Routledge.<\/p>\n<p class=\"import-Normal\">Huxley, Thomas. 1863. <em>Evidence as to Man's Place in Nature<\/em>. London: Williams &amp; Norgate.<\/p>\n<p class=\"import-Normal\">Jablonka, Eva, and Marion J. Lamb. 2005. <em>Evolution in Four Dimensions: Genetic, Epigenetic, Behavioral, and Symbolic Variation in the History of Life<\/em>. Cambridge, MA: The MIT Press.<\/p>\n<p class=\"import-Normal\">Kuklick, Henrika, ed. 2008. <em>A New History of Anthropology<\/em>. New York: Blackwell.<\/p>\n<p class=\"import-Normal\">Laland, Kevin N., Tobias Uller, Marcus W. Feldman, Kim Sterelny, Gerd B. Muller, Armin Moczek, Eva Jablonka, and John Odling-Smee. 2015. \u201cThe Extended Evolutionary Synthesis: Its Structure, Assumptions and Predictions.\u201d <em>Proceedings of the Royal Society, Series B<\/em> 282 (1813): 20151019.<\/p>\n<p class=\"import-Normal\">Lamarck, Jean Baptiste. 1809. <em>Philosophie Zoologique<\/em>. Paris: Dentu.<\/p>\n<p class=\"import-Normal\">Landau, Misia. 1991. <em>Narratives of Human Evolution<\/em>. New Haven: Yale University Press.<\/p>\n<p class=\"import-Normal\">Lee, Sang-Hee. 2017. <em>Close Encounters with Humankind: A Paleoanthropologist Investigates Our Evolving Species<\/em>. New York: W. W. Norton.<\/p>\n<p class=\"import-Normal\">Livingstone, David N. 2008. <em>Adam's Ancestors: Race, Religion, and the Politics of Human Origins<\/em>. Baltimore: Johns Hopkins University Press.<\/p>\n<p class=\"import-Normal\">Marks, Jonathan. 2015. <em>Tales of the Ex-Apes: How We Think about Human Evolution<\/em>. Berkeley, CA: University of California Press.<\/p>\n<p class=\"import-Normal\">Pigliucci, Massimo. 2009. \u201cThe Year in Evolutionary Biology 2009: An Extended Synthesis for Evolutionary Biology.\u201d <em>Annals of the New York Academy of Sciences<\/em> 1168: 218\u2013228.<\/p>\n<p class=\"import-Normal\">Simpson, George Gaylord. 1949. <em>The Meaning of Evolution: A Study of the History of Life and of Its Significance for Man<\/em>. New Haven: Yale University Press.<\/p>\n<p class=\"import-Normal\">Sommer, Marianne. 2016.<em> History Within: The Science, Culture, and Politics of Bones, Organisms, and Molecules<\/em>. Chicago: University of Chicago Press.<\/p>\n<p class=\"import-Normal\">Stoczkowski, Wiktor. 2002. <em>Explaining Human Origins: Myth, Imagination and Conjecture<\/em>. New York: Cambridge University Press.<\/p>\n<p class=\"import-Normal\">Tattersall, Ian, and Rob DeSalle. 2019. <em>The Accidental Homo sapiens: Genetics, Behavior, and Free Will<\/em>. New York: Pegasus.<\/p>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\">Barton, Robert A. 1996. \"Neocortex Size and Behavioural Ecology in Primates.\" <em>Proceedings of the Royal Society of London. Series B: Biological Sciences<\/em> 263 (1367): 173\u2013177.<\/p>\n<p class=\"import-Normal\">Bodmer, Walter, and Robin McKie. 1997. <em>The Book of Man: The Hman Genome Project and the Quest to Discover our Genetic Heritage.<\/em> Oxford University Press.<\/p>\n<p class=\"import-Normal\">Darwin, Charles. 1859.<em> On the Origin of Species by Means of Natural Selection, or, the Preservation of Favoured Races in the Struggle for Life<\/em>. London: J. Murray.<\/p>\n<p class=\"import-Normal\">Darwin, Charles. 1871. <em>The Descent of Man, and Selection in Relation to Sex.<\/em> London: J. Murray.<\/p>\n<p class=\"import-Normal\">Dawkins, Richard. 1976. <em>The Selfish Gene. <\/em>Oxford University Press.<\/p>\n<p class=\"import-Normal\">Deacon, T. W. 1998. <em>The Symbolic Species: The Co-evolution of Language and the Brain<\/em>. W. W. Norton &amp; Company.<\/p>\n<p class=\"import-Normal\">Eldredge, N., and S. J. Gould. 1972. \"Punctuated Equilibria: An Alternative to Phyletic Gradualism.\" In <em>Models in Paleobiology<\/em>, edited by T. J. Schopf, 82\u2013115. San Francisco: W. H. Freeman.<\/p>\n<p class=\"import-Normal\">Gould, Stephen J. 2003.<em> The Structure of Evolutionary Theory<\/em>. Cambridge, MA: Harvard University Press.<\/p>\n<p class=\"import-Normal\">Gould, Stephen J. 1996. <em>Mismeasure of Man<\/em>. New York: WW Norton &amp; Company.<\/p>\n<p class=\"import-Normal\">Gould, Stephen Jay, and Richard C. Lewontin. 1979. \"The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme.\" <em>Proceedings of the Royal Society of London. Series B: Biological Sciences<\/em> 205 (1151): 581\u2013598.<\/p>\n<p class=\"import-Normal\">Haeckel, Ernst. 1868. <em>Nat\u00fcrliche Sch\u00f6pfungsgeschichte<\/em>. Berlin: Reimer.<\/p>\n<p class=\"import-Normal\">Huxley, Thomas Henry. 1863. <em>Evidence as to Man\u2019s Place in Nature. <\/em>London: Williams and Norgate.<\/p>\n<p class=\"import-Normal\">Kaufman, Thomas C., Mark A. Seeger, and Gary Olsen. 1990. \"Molecular and Genetic Organization of the Antennapedia Gene Complex of <em>Drosophila melanogaster<\/em>.\" <em>Advances in Genetics<\/em> 27: 309\u2013362.<\/p>\n<p class=\"import-Normal\">Kellogg, Vernon. 1917. <em>Headquarters Nights<\/em>. Boston: The Atlantic Monthly Press.<\/p>\n<p class=\"import-Normal\">Kevles, Daniel J., and Leroy Hood. 1993. <em>The Code of Codes: Scientific and Social Issues in the Human Genome Project<\/em>. Cambridge, MA: Harvard University Press.<\/p>\n<p class=\"import-Normal\">Lewontin, Richard, Steven Rose, and Leon Kamin. 2017. <em>Not in Our Genes\u202f: Biology, Ideology, and Human Nature<\/em>, 2nd ed. Chicago: Haymarket Books.<\/p>\n<p class=\"import-Normal\">Lloyd, Elisabeth A., and Stephen J. Gould. 1993. \"Species Selection on Variability.\" <em>Proceedings of the National Academy of Sciences<\/em> 90 (2): 595\u2013599.<\/p>\n<p class=\"import-Normal\">Marks, Jonathan. 2015. \u201cThe Biological Myth of Human Evolution.\u201d In <em>Biologising the Social Sciences: Challenging Darwinian and Neuroscience Explanations<\/em>, edited by David Canter and David A. Turner, 59\u201378. London: Routledge.<\/p>\n<p class=\"import-Normal\">Monypenny, William Flavelle, and George Earle Buckle. 1929. <em>The Life of Benjamin Disraeli, Earl of Beaconsfield, Volume II: 1860\u20131881<\/em>. London: John Murray.<\/p>\n<p class=\"import-Normal\">Potts, Rick. 1998. \u201cVariability Selection in Hominid Evolution.\u201d <em>Evolutionary Anthropology <\/em><em>7<\/em><em>:<\/em> 81\u201396.<\/p>\n<p class=\"import-Normal\">Punnett, R. C. 1905. <em>Mendelism<\/em>. Cambridge: Macmillan and Bowes.<\/p>\n<p class=\"import-Normal\">Shapiro, Robert. 1991. <em>The Human Blueprint: The Race to Unlock the Secrets of Our Genetic Script.<\/em> New York: St. Martin\u2019s Press.<\/p>\n<p class=\"import-Normal\">Shultz, Susanne, Emma Nelson, and Robin Dunbar. 2012. \"Hominin Cognitive Evolution: Identifying Patterns and Processes in the Fossil and Archaeological Record.\" <em>Philosophical Transactions of the Royal Society B: Biological Sciences<\/em> 367 (1599): 2130\u20132140.<\/p>\n<p class=\"import-Normal\">Spencer, Herbert. 1864. <em>Principles of Biology.<\/em> London: Williams and Norgate.<\/p>\n<p class=\"import-Normal\">Watson, James D. 1990. \"The Human Genome Project: Past, Present, and Future.\" <em>Science<\/em> 248 (4951): 44\u201349.<\/p>\n<p class=\"import-Normal\">Yengo, L., Vedantam, S., Marouli, E., Sidorenko, J., Bartell, E., Sakaue, S., Graff, M., Eliasen, A.U., Jiang, Y., Raghavan, S. and Miao, J., 2022. A saturated map of common genetic variants associated with human height. <em>Nature<\/em>, <em>610 <\/em>(7933): 704-712.<\/p>\n<p class=\"import-Normal\">Zeder, Melinda A. 2018. \"Why Evolutionary Biology Needs Anthropology: Evaluating Core Assumptions of the Extended Evolutionary Synthesis.\" <em>Evolutionary Anthropology: Issues, News, and Reviews<\/em> 27 (6): 267\u2013284.<\/p>\n<\/div>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1763\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1763\"><div tabindex=\"-1\"><p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kerryn Warren, Ph.D., Grad Coach International<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lindsay Hunter, M.A., University of Iowa<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Navashni Naidoo, M.Sc., University of Cape Town<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Silindokuhle Mavuso, M.Sc., University of Witwatersrand<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\"><em>This chapter is a revision from <\/em><em>\"<\/em><a class=\"rId7\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\"><em>Chapter 9: Early Hominins<\/em><\/a><em>\" <\/em><em>by Kerryn Warren, K. Lindsay Hunter, Navashni Naidoo, Silindokuhle Mavuso, Kimberleigh Tommy, Rosa Moll, and Nomawethu Hlazo<\/em><em>. In <\/em><a class=\"rId8\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, first edition<\/em><\/a><em>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under <\/em><a class=\"rId9\" style=\"color: #000000\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\"><em>CC BY-NC 4.0<\/em><\/a><em>. <\/em><\/span><\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\"><span style=\"color: #000000\"><span style=\"color: #ffffff\">Learning Objectives<\/span><br \/>\n<\/span><\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li><span style=\"color: #000000\">Understand what is meant by \u201cderived\u201d and \u201cancestral\u201d traits and why this is relevant for understanding early hominin evolution.<\/span><\/li>\n<li><span style=\"color: #000000\">Understand changing paleoclimates and paleoenvironments as potential factors influencing early hominin adaptations.<\/span><\/li>\n<li><span style=\"color: #000000\">Describe the anatomical changes associated with bipedalism and dentition in early hominins, as well as their implications..<\/span><\/li>\n<li><span style=\"color: #000000\">Describe early hominin genera and species, including their currently understood dates and geographic expanses.<\/span><\/li>\n<li><span style=\"color: #000000\">Describe the earliest stone tool techno-complexes and their impact on the transition from early hominins to our genus.<\/span><\/li>\n<\/ul>\n<\/div>\n<\/div>\n<h2 class=\"__UNKNOWN__\"><span style=\"color: #000000\">Defining Hominins<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">It is through our study of our hominin ancestors and relatives that we are exposed to a world of \u201cmight have beens\u201d: of other paths not taken by our species, other ways of being human. But to better understand these different evolutionary trajectories, we must first define the terms we are using. If an imaginary line were drawn between ourselves and our closest relatives, the great apes, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1200\">bipedalism<\/a><\/strong> (or habitually walking upright on two feet) is where that line would be. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1270\">Hominin<\/a><\/strong>, then, means everyone on \u201cour\u201d side of the line: humans and all of our extinct bipedal ancestors and relatives since our divergence from the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1272\">last common ancestor (LCA)<\/a><\/strong> we share with chimpanzees.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Historic interpretations of our evolution, prior to our finding of early hominin <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1274\"><strong>fossils<\/strong><\/a>, varied. Debates in the mid-1800s regarding hominin origins focused on two key issues:<\/span><\/p>\n<ul>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">Where did we evolve?<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">Which traits evolved first?<\/span><\/li>\n<\/ul>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff00ff\">Charles Darwin hypothesized that we evolved in Africa, as he was convinced that we shared greater commonality with chimpanzees and gorillas on the continent (Darwin 1871). Others, such as Ernst Haeckel and Eug\u00e8ne Dubois, insisted that we were closer in affinity to orangutans and that we evolved in Eurasia where, until the discovery of the Taung Child in South Africa in 1924, all humanlike fossils (of Neanderthals and <em>Homo erectus<\/em>) had been found (Shipman 2002).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Within this conversation, naturalists and early <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1276\">paleoanthropologists<\/a><\/strong> (people who study human evolution) speculated about which human traits came first. These included the evolution of a big brain (<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1278\"><strong>encephalization<\/strong><\/a>), the evolution of the way in which we move about on two legs (bipedalism), and the evolution of our flat faces and small teeth (indications of dietary change). Original hypotheses suggested that, in order to be motivated to change diet and move about in a bipedal fashion, the large brain needed to have evolved first, as is seen in the fossil species mentioned above.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">However, we now know that bipedal locomotion is one of the first things that evolved in our lineage, with early relatives having more apelike dentition and small brain sizes. While brain size expansion is seen primarily in our genus, <em>Homo<\/em>, earlier hominin brain sizes were highly variable between and within taxa, from 300 cc (cranial capacity, cm<sup>3<\/sup>), estimated in <em>Ardipithecus<\/em>, to 550 cc, estimated in <em>Paranthropus boisei<\/em>. The lower estimates are well within the range of variation of nonhuman extant great apes. In addition, body size variability also plays a role in the interpretation of whether brain size could be considered large or small for a particular species or specimen. In this chapter, we will tease out the details of early hominin evolution in terms of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1280\"><strong>morphology<\/strong><\/a> (i.e. the study of the form, size, or shape of things; in this case, skeletal parts).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">We also know that early human evolution occurred in a very complicated fashion. There were multiple species (multiple genera) that featured diversity in their diets and locomotion. Specimens have been found all along the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1282\"><strong>East African Rift System <\/strong>(<strong>EARS)<\/strong><\/a>; that is, in Ethiopia, Kenya, Tanzania, and Malawi; see Figure 9.1), in limestone caves in South Africa, and in Chad. Dates of these early relatives range from around 7 million years ago (mya) to around 1 mya, overlapping temporally with members of our genus, <em>Homo<\/em>.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 610px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/06\/image38.png\" alt=\"Patchy green mountain alongside a deep sandy valley in East Africa.\" width=\"610\" height=\"277\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.1: East African Rift System (EARS). Credit: <a href=\"https:\/\/www.flickr.com\/photos\/ninara\/8624605781\/in\/photolist-x2yH7-x2yHe-VfVWuD-e98mPF-SzzjsU-2bsBZhC-2hHec7m-xtJ7Ez-NXnXvh-7Yg3uo-2cS3FgG-2hjo1Dc-2hjGoTS-nnumi8-82U66W-dMNn7B-8jdVbd-NWDg8-NW6fj-ebhx5w-bkFv1G-Ct5ZD-5JQk8A-y6TgAc-x9k6oe-2ebLTDC-WcPMnJ-2ekh6CS-Cu3LH-xNHDFK-9RUsZi-94jVt4-P46uiB-QFyjyE-crU8N7-5JLJKV-2ekSgk8-5JL454-2cPgZrF-2bHfQZu-dMTVPN-6yUbeN-jzMicQ-48XjU9-2etR2Ze-Styrvw-crU7V7-2wakq3-crU6Z1-2etR2XR\/\">IMG_1696 Great Rift Valley<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/ninara\/\">Ninara<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\">CC BY 2.0 License<\/a>.<\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Yet there is still so much to understand. Modern debates now look at the relatedness of these species to us and to one another, and they consider which of these species were able to make and use tools. As a result, every <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1284\">site<\/a><\/strong> discovery in the patchy hominin fossil record tells us more about our evolution. In addition, recent scientific techniques (not available even ten years ago) provide new insights into the diets, environments, and lifestyles of these ancient relatives.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In the past, <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1288\"><strong>taxonom<\/strong><strong>y<\/strong><\/a> was primarily based on morphology. Today it is tied to known relationships based on molecular <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1290\"><strong>phylogeny<\/strong><\/a> (e.g., based on DNA) or a combination of the two. This is complicated when applied to living <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1286\"><strong>taxa<\/strong><\/a>, but becomes much more difficult when we try to categorize ancestor-descendant relationships for long-extinct species whose molecular information is no longer preserved. We therefore find ourselves falling back on morphological comparisons, often of teeth and partially fossilized skeletal material.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">It is here that we turn to the related concepts of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1292\">cladistics<\/a> <\/strong>and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1294\">phylogenetics<\/a><\/strong><strong>. <\/strong><span style=\"background-color: #ff99cc\">Cladistics groups organisms according to their last common ancestors based on shared <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1126\">derived traits<\/a><\/strong>. <\/span>In the case of early hominins, these are often morphological traits that differ from those seen in earlier populations. These new or modified traits provide evidence of evolutionary relationships, and organisms with the same derived traits are grouped in the same <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1116\">clade<\/a> <\/strong>(Figure 9.2). <span style=\"background-color: #ff99cc\">For example, if we use feathers as a trait, we can group pigeons and ostriches into the clade of birds.<\/span> In this chapter, we will examine the grouping of the Robust Australopithecines, whose cranial and dental features differ from those of earlier hominins, and therefore are considered derived.<\/span><\/p>\n<figure style=\"width: 708px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image48.png\" alt=\"Phylogenetic tree shows clades and non clade groupings.\" width=\"708\" height=\"192\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.2: Clades refer to groups of species or taxa that share a common ancestor. In <span class=\"ILfuVd\" lang=\"en\"><span class=\"hgKElc\">a phylogeny, a clade is a complete group of lineages, including their last common ancestor. Groupings that do not include a common ancestor and <em>all<\/em> of its descendants are not clades. <\/span><\/span>Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\">Clades (Figure 9.2)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Katie Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Dig Deeper: Problems Defining Hominin Species<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000\">It is worth noting that species designations for early hominin specimens are often highly contested. This is due to the fragmentary nature of the fossil record, the large timescale (millions of years) with which paleoanthropologists need to work, and the difficulty in evaluating whether morphological differences and similarities are due to meaningful phylogenetic or biological differences or subtle differences\/variation in niche occupation or time. In other words, do morphological differences really indicate different species? How would classifying species in the paleoanthropological record compare with classifying living species today, for whom we can sequence genomes and observe lifestyles?<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">There are also broader philosophical differences among researchers when it comes to paleo-species designations. Some scientists, known as \u201c<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1296\"><strong>lumpers<\/strong>,<\/a>\u201d argue that large variability is expected among multiple populations in a given species over time. These researchers will therefore prefer to \u201clump\u201d specimens of subtle differences into single taxa. Others, known as \u201c<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1298\"><strong>splitters<\/strong><\/a>,\u201d argue that species variability can be measured and that even subtle differences can imply differences in niche occupation that are extreme enough to mirror modern species differences. In general, splitters would consider geographic differences among populations as meaning that a species is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1300\">polytypic<\/a><\/strong> <span style=\"background-color: #ff99cc\">(i.e., capable of interacting and breeding biologically but having morphological population differences)<\/span>. This is worth keeping in mind when learning about why species designations may be contested.<\/span><\/p>\n<figure style=\"width: 520px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image8.jpg\" alt=\"A graph shows a curved line depicting changes in morphology among two species over time.\" width=\"520\" height=\"292\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.3: This graph demonstrates the concept of a chronospecies, where one species (Species A) \u201cevolves\u201d into another (Species B). Credit: Chronospecies original to Explorations: An Open Invitation to Biological Anthropology, 2nd edition by Kerryn Warren is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\"><span style=\"color: #000000\">This further plays a role in evaluating ancestry. Debates over which species \u201cgave rise\u201d to which continue to this day. It is common to try to create \u201clineages\u201d of species to determine when one species evolved into another over time. We refer to these as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1302\"><strong>chronospecies<\/strong><\/a> (Figure 9.3). Constructed hominin phylogenetic trees are routinely variable, changing with new specimen discoveries, new techniques for evaluating and comparing species, and, some have argued, nationalist or biased interpretations of the record. More recently, some researchers have shifted away from \u201ctreelike\u201d models of ancestry toward more nuanced metaphors such as the \u201cbraided stream,\u201d where some levels of interbreeding among species and populations are seen as natural processes of evolution.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Finally, it is worth considering the process of fossil discovery and publication. Some fossils are easily diagnostic to a species level and allow for easy and accurate interpretation. Some, however, are more controversial. This could be because they do not easily preserve or are incomplete, making it difficult to compare and place within a specific species (e.g., a fossil of a patella or knee bone). Researchers often need to make several important claims when announcing or publishing a find: a secure date (if possible), clear association with other finds, and an adequate comparison among multiple species (both extant and fossil). Therefore, it is not uncommon that an important find was made years before it is scientifically published.<\/span><\/p>\n<\/div>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Paleoenvironment and Hominin Evolution<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There is no doubt that one of the major selective pressures in hominin evolution is the environment. <span style=\"background-color: #ff9900\"><span style=\"background-color: #ffff00\">Large-scale changes in global and regional climate, as well as alterations to the environment, are<\/span><span style=\"background-color: #ffff00\"> (<\/span><\/span><em style=\"background-color: transparent\"><span style=\"text-decoration: underline\">thought to be<\/span>)<\/em><span style=\"background-color: #ffff00\"><del> all<\/del> linked to (all) hominin diversification, dispersal, and extinction<\/span> (Maslin et al. 2014). Environmental reconstructions often use modern analogues. Let us take, for instance, the hippopotamus. It is an animal that thrives in environments that have abundant water to keep its skin cool and moist. If the environment for some reason becomes drier, it is expected that hippopotamus populations will reduce. If a drier environment becomes wetter, it is possible that hippopotamus populations may be attracted to the new environment and thrive. Such instances have occurred multiple times in the past, and the bones of some <strong style=\"background-color: transparent\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1308\">fauna<\/a><\/strong> (i.e., animals, like the hippopotamus) that are sensitive to these changes give us insights into these events.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Yet reconstructing a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1304\">paleoenvironment<\/a><\/strong> relies on a range of techniques, which vary depending on whether research interests focus on local changes or more global environmental changes\/reconstructions. For local environments (such as a single site or region), comparing the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1306\">faunal assemblages<\/a> <\/strong>(collections of fossils of animals found at a site) with animals found in certain modern environments allows us to determine if past environments mirror current ones in the region. Changes in the faunal assemblages, as well as when they occur and how they occur, tell us about past environmental changes. Other techniques are also useful in this regard. Chemical analyses, for instance, can reveal the diets of individual fauna, providing clues as to the relative wetness or dryness of their environment (e.g., nitrogen <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1310\"><strong>isotopes<\/strong><\/a>; Kingston and Harrison 2007).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Global climatic changes in the distant past, which fluctuated between being colder and drier and warmer and wetter on average, would have global implications for environmental change (Figure 9.4). These can be studied by comparing marine core and terrestrial soil data across multiple sites. These techniques are based on chemical analysis, such as examination of the nitrogen and oxygen isotopes in shells and sediments. Similarly, analyzing pollen grains shows which kinds of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1312\"><strong>flora<\/strong> <\/a> survived in an environment at a specific time period. There are multiple lines of evidence that allow us to visualize global climate trends over millions of years (although it should be noted that the direction and extent of these changes could differ by geographic region).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 649px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image12-1-1.png\" alt=\"Chart shows cyclical carbon dioxide levels from 800,000 years ago until today.\" width=\"649\" height=\"406\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.4: This graph, based on the comparison of atmospheric samples contained in ice cores and more recent direct measurements, illustrates how atmospheric CO\u2082 has fluctuated over time and increased sharply since the Industrial Revolution. The graph also shows that since 800,000ya (and before) atmospheric CO\u2082 has never exceeded 300 parts per million (ppm). In 1950 it was 310ppm. Today atmospheric CO\u2082 has spiked to over 410 ppm. Credit: <a href=\"https:\/\/climate.nasa.gov\/evidence\/\">CO\u2082 increase since the Industrial Revolution<\/a> by <a href=\"https:\/\/www.nasa.gov\/\">NASA<\/a> is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a> and is used within <a href=\"https:\/\/www.nasa.gov\/multimedia\/guidelines\/index.html\">NASA guidelines on re-use<\/a>. Original from Luthi, D., et al.. 2008; Etheridge, D.M., et al. 2010; Vostok ice core data\/J.R. Petit et al.; NOAA Mauna Loa CO<a href=\"https:\/\/climate.nasa.gov\/evidence\/\">\u2082<\/a> record..<\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Both local and global climatic\/environmental changes have been used to understand factors affecting our evolution (DeHeinzelin et al. 1999; Kingston 2007). Environmental change acts as an important factor regarding the onset of several important hominin traits seen in early hominins and discussed in this chapter. Namely, the environment has been interpreted as the following:<\/span><\/p>\n<ul>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">the driving force behind the evolution of bipedalism,<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">the reason for change and variation in early hominin diets, and<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">the diversification of multiple early hominin species.<\/span><\/li>\n<\/ul>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are numerous hypotheses regarding how climate has driven and continues to drive human evolution. Here, we will focus on just three popular hypotheses.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Savannah Hypothesis (or Aridity Hypothesis)<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The hypothesis:<\/strong> This popular theory suggests that the expansion of the savannah (or less densely forested, drier environments) forced early hominins from an <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1314\"><strong>arboreal<\/strong> <\/a> lifestyle (one living in trees) to a terrestrial one where bipedalism was a more efficient form of locomotion (Figure 9.5). It was first proposed by Darwin (1871) and supported by anthropologists like Raymond Dart (1925). However, this idea was supported by little fossil or paleoenvironmental evidence and was later refined as the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1318\">Aridity Hypothesis<\/a><\/strong>. This hypothesis states that the long-term <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1316\">aridification<\/a><\/strong> and, thereby, expansion of savannah biomes were drivers in diversification in early hominin evolution (deMenocal 2004; deMenocal and Bloemendal 1995). It advocates for periods of accelerated aridification leading to early hominin speciation events.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 647px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image46.png\" alt=\"Photograph showing a dry, open savannah environment.\" width=\"647\" height=\"486\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.5: The African savannah grew during early hominin evolution. This may have forced early hominins from an arboreal lifestyle to a terrestrial one, where bipedalism was a more efficient form of locomotion. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:African_savannah_@_Masai_Mara_(21308330314).jpg\">African savannah @ Masai Mara (21308330314)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/132394214@N04\">Leo Li<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\">CC BY 2.0 License<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The evidence:<\/strong> While early bipedal hominins are often associated with wetter, more closed environments (i.e., not the Savannah Hypothesis), both marine and terrestrial records seem to support general cooling, drying conditions, with isotopic records indicating an increase in grasslands (i.e., colder and wetter climatic conditions) between 8 mya and 6 mya across the African continent (Cerling et al. 2011). This can be contrasted with later climatic changes derived from aeolian dust records (sediments transported to the site of interest by wind), which demonstrate increases in seasonal rainfall between 3 mya and 2.6 mya, 1.8 mya and 1.6 mya, and 1.2 mya and 0.8 mya (deMenocal 2004; deMenocal and Bloemendal 1995).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Interpretation(s):<\/strong> Despite a relatively scarce early hominin record, it is clear that two important factors occur around the time period in which we see increasing aridity. The first factor is the diversification of taxa, where high morphological variation between specimens has led to the naming of multiple hominin genera and species. The second factor is the observation that the earliest hominin fossils appear to have traits associated with bipedalism and are dated to around the drying period (as based on isotopic records). Some have argued that it is more accurately a combination of bipedalism and arboreal locomotion, which will be discussed later. However, the local environments in which these early specimens are found (as based on the faunal assemblages) do not appear to have been dry.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Turnover Pulse Hypothesis<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The hypothesis:<\/strong> In 1985, paleontologist Elisabeth Vbra noticed that in periods of extreme and rapid climate change, <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1326\"><strong>ungulates<\/strong><\/a> (hoofed mammals of various kinds) that had generalized diets fared better than those with specialized diets (Vrba 1988, 1998). <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1322\"><strong>Specialist<\/strong><\/a> eaters <span style=\"background-color: #ff99cc\">(those who rely primarily on specific food types)<\/span> faced extinction at greater rates than their <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1320\">generalist<\/a> <\/strong><span style=\"background-color: #ff99cc\">(those who can eat more varied and variable diets) <\/span>counterparts because they were unable to adapt to new environments (Vrba 2000). Thus, periods with extreme climate change would be associated with high <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1324\"><strong>faunal turnover<\/strong><\/a>: that is, the extinction of many species and the speciation, diversification, and migration of many others to occupy various niches.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The evidence:<\/strong> The onset of the<strong> <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1400\">Quaternary Ice Age<\/a><\/strong>, between 2.5 mya and 3 mya, brought extreme global, cyclical <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1328\"><strong>interglacial<\/strong> <\/a> and <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1330\"><strong>glacial<\/strong><\/a> periods (warmer, wetter periods with less ice at the poles, and colder, drier periods with more ice near the poles). Faunal evidence from the Turkana basin in East Africa indicates multiple instances of faunal turnover and extinction events, in which global climatic change resulted in changes from closed\/forested to open\/grassier habitats at single sites (Behrensmeyer et al. 1997; Bobe and Behrensmeyer 2004). Similarly, work in the Cape Floristic Belt of South Africa shows that extreme changes in climate play a role in extinction and migration in ungulates. While this theory was originally developed for ungulates, its proponents have argued that it can be applied to hominins as well. However, the link between climate and speciation is only vaguely understood (Faith and Behrensmeyer 2013).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Interpretation(s):<\/strong> While the evidence of rapid faunal turnover among ungulates during this time period appears clear, there is still some debate around its usefulness as applied to the paleoanthropological record. Specialist hominin species do appear to exist for long periods of time during this time period, yet it is also true that <em>Homo<\/em>, a generalist genus with a varied and adaptable diet, ultimately survives the majority of these fluctuations, and the specialists appear to go extinct.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Variability Selection Hypothesis<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The hypothesis: <\/strong>This hypothesis was first articulated by paleoanthropologist Richard Potts (1998). It links the high amount of climatic variability over the last 7 million years to both behavioral and morphological changes. Unlike previous notions, this hypothesis states that hominin evolution does not respond to habitat-specific changes or to specific aridity or moisture trends. Instead, long-term environmental unpredictability over time and space influenced morphological and behavioral adaptations that would help hominins survive, regardless of environmental context (Potts 1998, 2013). The Variability Selection Hypothesis states that hominin groups would experience varying degrees of natural selection due to continually changing environments and potential group isolation. This would allow certain groups to develop genetic combinations that would increase their ability to survive in shifting environments. These populations would then have a genetic advantage over others that were forced into habitat-specific adaptations (Potts 2013).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The evidence:<\/strong> The evidence for this theory is similar to that for the Turnover Pulse Hypothesis: large climatic variability and higher survivability of generalists versus specialists. However, this hypothesis accommodates for larger time-scales of extinction and survival events.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Interpretation(s):<\/strong> In this way, the Variability Selection Hypothesis allows for a more flexible interpretation of the evolution of bipedalism in hominins and a more fluid interpretation of the Turnover Pulse Hypothesis, where species turnover is meant to be more rapid. In some ways, this hypothesis accommodates both environmental data and our interpretations of an evolution toward greater variability among species and the survivability of generalists.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000;background-color: #ff99cc\"><strong>Paleoenvironment Summary<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff99cc\">Some hypotheses presented in this section pay specific attention to habitat (Savannah Hypothesis) while others point to large-scale climatic forces (Variability Selection Hypothesis). Some may be interpreted to describe the evolution of traits such as bipedalism (Savannah Hypothesis), and others generally explain the diversification of early hominins (Turnover Pulse and Variability Selection Hypotheses). While there is no consensus as to how the environment drove our evolution, it is clear that the environment shaped both habitat and resource availability in ways that would have influenced our early ancestors physically and behaviorally.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Derived Adaptations: Bipedalism<br \/>\n<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The unique form of locomotion exhibited by modern humans, called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1332\">obligate bipedalism<\/a><\/strong>, is important in distinguishing our species from the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1334\"><strong>extant<\/strong><\/a> (living) great apes. The ability to walk habitually upright is thus considered one of the defining attributes of the hominin lineage. We also differ from other animals that walk bipedally (such as kangaroos) in that we do not have a tail to balance us as we move.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The origin of bipedalism in hominins has been debated in paleoanthropology, but at present there are two main <span style=\"background-color: #ff99cc\">ideas<\/span>: <span style=\"text-decoration: underline\">(<em>theories<\/em>)<\/span><\/span><\/p>\n<ol>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">early hominins initially lived in trees, but increasingly started living on the ground, so we were a product of an arboreal last common ancestor (LCA) or,<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">our LCA was a terrestrial quadrupedal knuckle-walking species, more similar to extant chimpanzees.<\/span><\/li>\n<\/ol>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Most research supports the first theory of an arboreal LCA based on skeletal morphology of early hominin genera that demonstrate adaptations for climbing but not for knuckle-walking. This would mean that both humans and chimpanzees can be considered \u201cderived\u201d in terms of locomotion since chimpanzees would have independently evolved knuckle-walking.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are many current ideas regarding selective pressures that would lead to early hominins adapting upright posture and locomotion. Many of these selective pressures, as we have seen in the previous section, coincide with a shift in environmental conditions, supported by paleoenvironmental data. In general, however, it appears that, like extant great apes, early hominins thrived in forested regions with dense tree coverage, which would indicate an arboreal lifestyle. As the environmental conditions changed and a savannah\/grassland environment became more widespread, the tree cover would become less dense, scattered, and sparse such that bipedalism would become more important.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are several proposed selective pressures for bipedalism:<\/span><\/p>\n<ol>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\"><strong>E<\/strong><strong>nergy conservation:<\/strong> Modern bipedal humans conserve more energy than extant chimpanzees, which are predominantly knuckle-walking quadrupeds when walking over land. While chimpanzees, for instance, are faster than humans terrestrially, they expend large amounts of energy being so. Adaptations to bipedalism include \u201cstacking\u201d the majority of the weight of the body over a small area around the center of gravity (i.e., the head is above the chest, which is above the pelvis, which is over the knees, which are above the feet). This reduces the amount of muscle needed to be engaged during locomotion to \u201cpull us up\u201d and allows us to travel longer distances expending far less energy.<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\"><strong>T<\/strong><strong>hermoregulation:<\/strong> Less surface area (i.e., only the head and shoulders) is exposed to direct sunlight during the hottest parts of the day (i.e., midday). This means that the body has less need to employ additional \u201ccooling\u201d mechanisms such as sweating, which additionally means less water loss.<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Bipedalism <\/strong><span style=\"text-decoration: underline\">(Freeing of Hands)<\/span><strong>: <\/strong>This method of locomotion freed up our ancestors\u2019 hands such that they could more easily gather food and carry tools or infants. This further enabled the use of hands for more specialized adaptations associated with the manufacturing and use of tools.<\/span><\/li>\n<\/ol>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">These selective pressures are not mutually exclusive. Bipedality could have evolved from a combination of these selective pressures, in ways that increased the chances of early hominin survival.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Skeletal Adaptations for Bipedalism<\/strong><\/span><\/h3>\n<figure style=\"width: 405px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image10-1.png\" alt=\"A full human skeleton and gorilla skeleton standing in upright positions next to each other.\" width=\"405\" height=\"452\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.6: Compared to gorillas (right) and other apes, humans (left) have highly specialized adaptations to facilitate bipedal locomotion. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Primatenskelett-drawing.jpg\">Skeleton of human (1) and gorilla (2), unnaturally sketched<\/a> by unknown from Brehms Tierleben, Small Edition 1927 is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Humans have highly specialized adaptations to facilitate obligate bipedalism (Figure 9.6). Many of these adaptations occur within the soft tissue of the body (e.g., muscles and tendons). However, when analyzing the paleoanthropological record for evidence of the emergence of bipedalism, all that remains is the fossilized bone. Interpretations of locomotion are therefore often based on comparative analyses between fossil remains and the skeletons of extant primates with known locomotor behaviors. These adaptations occur throughout the skeleton and are summarized in Figure 9.7.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The majority of these adaptations occur in the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1336\"><strong>postcranium<\/strong><\/a> <span style=\"background-color: #ff99cc\">(the skeleton from below the head)<\/span> and are outlined in Figure 9.7. In general, these adaptations allow for greater stability and strength in the lower limb, by allowing for more shock absorption, for a larger surface area for muscle attachment, and for the \u201cstacking\u201d of the skeleton directly over the center of gravity to reduce energy needed to be kept upright. These adaptations often mean less flexibility in areas such as the knee and foot.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">However, these adaptations come at a cost. Evolving from a nonobligate bipedal ancestor means that the adaptations we have are evolutionary compromises. For instance, the valgus knee (angle at the knee) is an essential adaptation to balance the body weight above the ankle during bipedal locomotion. However, the strain and shock absorption at an angled knee eventually takes its toll. For example, runners often experience joint pain. Similarly, the long neck of the femur absorbs stress and accommodates for a larger pelvis, but it is a weak point, resulting in hip replacements being commonplace among the elderly, especially in cases where the bone additionally weakens through osteoporosis. Finally, the S-shaped curve in our spine allows us to stand upright, relative to the more curved C-shaped spine of an LCA. Yet the weaknesses in the curves can lead to pinching of nerves and back pain. Since many of these problems primarily are only seen in old age, they can potentially be seen as an evolutionary compromise.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Despite relatively few postcranial fragments, the fossil record in early hominins indicates a complex pattern of emergence of bipedalism. Key features, such as a more anteriorly placed foramen magnum, are argued to be seen even in the earliest discovered hominins, indicating an upright posture (Dart 1925). Some early species appear to have a mix of ancestral (arboreal) and derived (bipedal) traits, which indicates a mixed locomotion and a more <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1402\">mosaic evolution<\/a><\/strong> of the trait. Some early hominins appear to, for instance, have bowl-shaped pelvises (hip bones) and angled femurs suitable for bipedalism but also have retained an opposable <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1338\">hallux<\/a><\/strong> (big toe) or curved fingers and longer arms (for arboreal locomotion). These mixed morphologies may indicate that earlier hominins were not fully obligate bipeds and thus thrived in mosaic environments.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Yet the associations between postcranial and the more diagnostic cranial fossils and bones are not always clear, muddying our understanding of the specific species to which fossils belong (Grine et al. 2022).<\/span><\/p>\n<div align=\"left\">\n<table class=\"grid\">\n<caption>\n<p class=\"import-Normal\" style=\"text-align: left\"><span style=\"color: #000000\">Figure 9.7: Skeletal comparisons between modern humans (obligate bipeds) and nonobligate bipeds (e.g., chimpanzees). Credit: <a class=\"rId34\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\">Skeletal comparisons between modern humans and <\/a><a class=\"rId35\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\">nonobligate<\/a><a class=\"rId36\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\"> bipeds (Figure 9.6)<\/a> original to <a class=\"rId37\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a class=\"rId38\" style=\"color: #000000\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/span><\/p>\n<\/caption>\n<thead>\n<tr>\n<td style=\"width: 97.998px\"><strong>Region<\/strong><\/td>\n<td style=\"width: 106.992px\"><strong>Feature<\/strong><\/td>\n<td style=\"width: 366.992px\"><strong>Obligate Biped (H. sapiens)<\/strong><\/td>\n<td style=\"width: 310px\"><strong>Nonobligate Biped<\/strong><\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr>\n<td style=\"width: 97.998px\">Cranium<\/td>\n<td style=\"width: 106.992px\">Position of the foramen magnum<\/td>\n<td style=\"width: 366.992px\">Positioned inferiorly (immediately under the cranium) so that the head rests on top of the vertebral column for balance and support (head is perpendicular to the ground).<\/td>\n<td style=\"width: 310px\">Posteriorly positioned (to the back of the cranium). Head is positioned parallel to the ground.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Body proportions<\/td>\n<td style=\"width: 366.992px\">Shorter upper limb (not used for locomotion).<\/td>\n<td style=\"width: 310px\">Longer upper limbs (used for locomotion).<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Spinal curvature<\/td>\n<td style=\"width: 366.992px\">S-curve due to pressure exerted on the spine from bipedalism (lumbar lordosis).<\/td>\n<td style=\"width: 310px\">C-curve.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Vertebrae<\/td>\n<td style=\"width: 366.992px\">Robust lumbar (lower-back) vertebrae (for shock absorbance and weight bearing). Lower back is more flexible than that of apes as the hips and trunk swivel when walking (weight transmission).<\/td>\n<td style=\"width: 310px\">Gracile lumbar vertebrae compared to those of modern humans.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Pelvis<\/td>\n<td style=\"width: 366.992px\">Shorter, broader, bowl-shaped pelvis (for support); very robust. Broad sacrum with large sacroiliac joint surfaces.<\/td>\n<td style=\"width: 310px\">Longer, flatter, elongated ilia; more narrow and gracile; narrower sacrum; relatively smaller sacroiliac joint surface.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Lower limb<\/td>\n<td style=\"width: 366.992px\">In general, longer, more robust lower limbs and more stable, larger joints.<\/p>\n<ul>\n<li style=\"font-weight: 400\">Large femoral head and longer neck (absorbs more stress and increases the mechanical advantage).<\/li>\n<li style=\"font-weight: 400\">Valgus knee, in which the angle of the knee positions it over the ankle and keeps the center of gravity balanced over the stance leg during stride cycle (shock absorbance).<\/li>\n<li style=\"font-weight: 400\">Distal tibia (lower leg) of humans has a large medial malleolus for stability.<\/li>\n<\/ul>\n<\/td>\n<td style=\"width: 310px\">In general, smaller, more gracile limbs with more flexible joints.<\/p>\n<ul>\n<li style=\"font-weight: 400\">Femoral neck is smaller in comparison to modern humans and shorter.<\/li>\n<li style=\"font-weight: 400\">The legs bow outward, and there is no valgus angle of the knee (no \u201cknock knees\u201d).<\/li>\n<li style=\"font-weight: 400\">The distal tibia in chimpanzees is trapezoid (wider anteriorly) for climbing and allows more flexibility.<\/li>\n<\/ul>\n<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Foot<\/td>\n<td style=\"width: 366.992px\">Rigid, robust foot, without a midtarsal break.<\/p>\n<p>Nonopposable and large, robust big toe (for push off while walking) and large heel for shock absorbance.<\/td>\n<td style=\"width: 310px\">Flexible foot, midtarsal break present (which allows primates to lift their heels independently from their feet), opposable big toe for grasping.<\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<p class=\"import-Normal\"><span style=\"color: #000000;background-color: #ccffcc\">It is also worth noting that, while not directly related to bipedalism per se, other postcranial adaptations are evident in the hominin fossil record from some of the earlier hominins. For instance, the hand and finger morphologies of many of the earliest hominins indicate adaptations consistent with arboreality. These include longer hands, more curved metacarpals and phalanges (long bones in the hand and fingers, respectively), and a shorter, relatively weaker thumb. This allows for gripping onto curved surfaces during locomotion. The earliest hominins appear to have mixed morphologies for both bipedalism and arborealism. However, among Australopiths (members of the genus, Australopithecus), there are indications for greater reliance on bipedalism as the primary form of locomotion. Similarly, adaptations consistent with tool manufacture (shorter fingers and a longer, more robust thumb, in contrast to the features associated with arborealism) have been argued to appear before the genus <em>Homo<\/em>.<\/span><\/p>\n<div class=\"textbox shaded\">\n<p><span style=\"background-color: #00ffff\">(Special Topic with student projects: fear of snakes, cultural or biological? Biology, culture, and the fear of snakes, Snake Detection Theory)<\/span><\/p>\n<p>It is suggested that primates have three major predators: raptors, felines, and snakes; however, many studies show that of these carnivores, snakes were one of the first that mammals had to contend with alongside dinosaurs, as felines and raptors evolved at a much slower pace than their reptilian competition. Herpetologists trace the evolution of constricting snakes to about 100 million years ago, and by the time mammals arrived around 75 million years ago, constrictors were\u00a0 already well established as a formidable threat (Greene, 2017). \u00a0Both co-existed for millennia and each sustained selective pressures requiring them to evolve specific traits to survive. When venomous snakes eventually emerged 55 to 65 million years ago, they posed yet an additional threat to proto-primates as they required less distance for the predator to kill (2017). Alongside camouflage and silent movement techniques, it was the development of the snake\u2019s hollow fangs through which to deliver venom that was most transformative to primate evolution. As such, primates evolved their pre-conscious attention, and visual acuity to cope with this new threat; therefore, while snakes were adapting morphologically to feed themselves, they were unwittingly teaching proto-primates valuable lessons in predator detection and reacting appropriately in order to survive.\u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0In a 2009 Harvard University study, Lynne A. Isbell hypothesizes that envenoming snakes are linked to being directly responsible for the origins of the evolving complex brains and superior visual capacity in the lineage of anthropoids leading to humans (Isbell, 2009). Forward-facing eyes for binocular vision, depth perception, enhanced visual acuity, stereoscopic and trichromatic colour vision, all traits necessary for snake detection; and the quick motor responses from the primate\u2019s fight, flight, or freeze defence mechanism to circumvent a snake\u2019s squeeze or bite. Numerous laboratory studies show that humans and primates both sense and visually detect snakes more rapidly than other threatening stimuli (Van Le Et al., 2013). These experiments show that snakes elicited the strongest, fastest responses (Van Le Et al., 2013). This is known as \u2018Snake Detection Theory\u2019 and is the evolution of the primate\u2019s complex brain, visual acuity, and rapid motor responses towards snakes in its environment that are the adaptations needed to live successfully as arboreal beings. It is not fortuitous then, that primates that never coexisted with venomous snakes, such as lemurs in Madagascar, have less visual acuity, better olfaction and smaller brains. Within Isbell\u2019s work, a collaborative study by a group of neuroscientists tested this hypothesis and found that, indeed, there is higher neural firing and activity in multiple areas of the primate brain, notably in the pulvinar, a region\u00a0 responsible for visual attention and oculomotor behaviour (Isbell, L., 2009).\u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 \u00a0 Today, the fear of snakes is widespread in humans, often shown through avoidance and disgust. A study in <em data-start=\"197\" data-end=\"244\">The Journal of Ethnobiology and Ethnomedicine<\/em> notes that snakes are over-hunted and excluded from conservation efforts worldwide (Cer\u00edaco, 2012). While cultural factors shape our sentiments, instinct also plays a role\u2014such as the developed avoidance behaviors toward threats like snakes. This blend of instinct and cultural influence is not only seen in behavior but also deeply embedded in the stories we tell. Many cultures depict mythological snakes as harbingers of death or chaos. In the Bible, Satan becomes a snake to tempt Eve. Norse mythology features J\u00f6rmungandr, the world serpent who signals the apocalypse. Egyptian myth tells of Apophis, who battles the sun god Ra nightly. Mesoamerican lore includes Quetzalcoatl, a feathered serpent linked to water and death. Though sources vary, these myths consistently portray snakes as threats. As such, the widespread fear of snakes may reflect both evolutionary and cultural influences. Understood as an adaptive response inherited from primate ancestors\u2014who developed avoidance behaviors toward potentially dangerous stimuli\u2014and reinforced through myths and religious narratives, the enduring presence of snakes as potent figures of fear across human societies and primate groups highlights the complex intertwining of instinct and cultural meaning in shaping human behavior.<\/div>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Early Hominins: Sahelanthropus and Orrorin<\/strong><\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">We see evidence for bipedalism in some of the earliest fossil hominins, dated from within our estimates of our divergence from chimpanzees. These hominins, however, also indicate evidence for arboreal locomotion.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The earliest dated hominin find (between 6 mya and 7 mya, based on radiometric dating of volcanic tufts) has been argued to come from Chad and is named <strong><em>Sahelanthropus tchadensis<\/em> <\/strong>(Figure 9.8; Brunet et al. 1995). The initial discovery was made in 2001 by Ahounta Djimdoumalbaye and announced in <em>Nature<\/em> in 2002 by a team led by French paleontologist Michel Brunet. The find has a small cranial capacity (360 cc) and smaller canines than those in extant great apes, though they are larger and pointier than those in humans. <span style=\"background-color: #ffff00\">This implies strongly that, over evolutionary time, the need for display and dominance among males has reduced, as has our sexual dimorphism<\/span>. A short cranial base and a foramen magnum that is more humanlike in positioning have been argued to indicate upright walking.<\/span><\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 640px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-289\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.82.jpg\" alt=\"Four views of a beige-colored skull are seen against a black background.\" width=\"640\" height=\"640\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.8: Sahelanthropus tchadensis exhibits a set of derived features, including a long, low cranium; a small, ape-sized braincase; and relatively reduced prognathism. Credit: aa <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 anterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 posterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 inferior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; and d <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 lateral left view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Initially, the inclusion of <em>Sahelanthropus<\/em> in the hominin family was debated by researchers, since the evidence for bipedalism is based on cranial evidence alone, which is not as convincing as postcranial evidence. Yet, a femur (thigh bone) and ulnae (upper arm bones) thought to belong to <em>Sahelanthropus<\/em> was discovered in 2001 (although not published until 2022). These bones may support the idea that the hominin was in fact a terrestrial biped with arboreal capabilities and behaviors (Daver et al. 2022).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Orrorin tugenensis<\/em><\/strong> (Orrorin meaning \u201coriginal man\u201d), dated to between 6 mya and 5.7 mya, was discovered near Tugen Hills in Kenya in 2000. Smaller <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1340\">cheek teeth<\/a><\/strong> (molars and premolars) than those in even more recent hominins, thick enamel, and reduced, but apelike, canines characterize this species. This is the first species that clearly indicates adaptations for bipedal locomotion, with fragmentary leg, arm, and finger bones having been found but few cranial remains. One of the most important elements discovered was a proximal femur, BAR 1002'00. The femur is the thigh bone, and the proximal part is that which articulates with the pelvis; this is very important for studying posture and locomotion. This femur indicates that <em>Ororrin<\/em> was bipedal, and recent studies suggest that it walked in a similar way to later <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1404\">Pliocene<\/a><\/strong> hominins. Some have argued that features of the finger bones suggest potential tool-making capabilities, although many researchers argue that these features are also consistent with climbing.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Early Hominins: The Genus <em>Ardipithecus<\/em><\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Another genus, <em>Ardipithecus<\/em>, is argued to be represented by at least two species: <em>Ardipithecus (Ar.) ramidus <\/em>and <em>Ar. kadabba<\/em>.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Ardipithecus ramidus<\/em><\/strong> (\u201cramid\u201d means root in the Afar language) is currently the best-known of the earliest hominins (Figure 9.9). Unlike <em>Sahelanthropus<\/em> and<em> Orrorin<\/em>, this species has a large sample size of over 110 specimens from Aramis alone. Dated to 4.4 mya, <em>Ar. ramidus<\/em> was found in Ethiopia (in the Middle Awash region and in Gona). This species was announced in 1994 by American palaeoanthropologist Tim White, based on a partial female skeleton nicknamed \u201cArdi\u201d (ARA-VP-6\/500; White et al. 1994). Ardi demonstrates a mosaic of ancestral and derived characteristics in the postcrania. For instance, she had an opposable big toe (hallux), similar to chimpanzees (i.e., more ancestral), which could have aided in climbing trees effectively. However, the pelvis and hip show that she could walk upright (i.e., it is derived), supporting her hominin status. A small brain (300 cc to 350 cc), midfacial projection, and slight prognathism show retained ancestral cranial features, but the cheek bones are less flared and robust than in later hominins.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 706px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-1310\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.9-scaled-1.jpg\" alt=\"Skull cast and partial skeleton with photographs of some bones and line drawings of others.\" width=\"706\" height=\"453\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.9a and b: Researchers believe that Ardipithecus ramidus was able to walk upright, although not as efficiently as later humans. It possessed the musculature required for tree climbing, and while moving quadrupedally, it likely placed weight on the palms of the hands rather than on the knuckles. Credit: a. <a class=\"rId61\" href=\"https:\/\/boneclones.com\/product\/ardipithecus-ramidus-skull-BH-039\">Ardipithecus ramidus Skull<\/a> by <a class=\"rId62\" href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a> is used by permission and available here under a <a class=\"rId63\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>; b. <a class=\"rId64\" href=\"https:\/\/boneclones.com\/product\/ardipithecus-ramidus-skull-BH-039\">Artist\u2019s rendition of \u201cArdi\u201d skeleton<\/a> by <a class=\"rId65\" href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a> is used by permission and available here under a <a class=\"rId66\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong><em>Ardipithecus kadabba<\/em><\/strong> (the species name means \u201coldest ancestor\u201d in the Afar language) is known from localities on the western margin of the Middle Awash region, the same locality where <em>Ar. ramidus<\/em> has been found. Specimens include mandibular fragments and isolated teeth as well as a few postcranial elements from the Asa Koma (5.5 mya to 5.77 mya) and Kuseralee Members (5.2 mya), well-dated and understood (but temporally separate) volcanic layers in East Africa. This species was discovered in 1997 by paleoanthropologist Dr. Yohannes Haile-Selassie. Originally these specimens were referred to as a subspecies of <em>Ar. ramidus<\/em>. In 2002, six teeth were discovered at Asa Koma and the dental-wear patterns confirmed that this was a distinct species, named <em>Ar. kadabba,<\/em> in 2004. One of the postcranial remains recovered included a 5.2 million-year-old toe bone that demonstrated features that are associated with toeing off (pushing off the ground with the big toe leaving last) during walking, a characteristic unique to bipedal walkers. However, the toe bone was found in the Kuseralee Member, and therefore some doubt has been cast by researchers about its association with the teeth from the Asa Koma Member.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"background-color: #ff99cc\"><strong><span style=\"color: #000000\">Bipedal Trends in Early Hominins: Summary<\/span><\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff99cc\">Trends toward bipedalism are seen in our earliest hominin finds. However, many specimens also indicate retained capabilities for climbing. Trends include a larger, more robust hallux; a more compact foot, with an arch; a robust, long femur, angled at the knee; a robust tibia; a bowl-shaped pelvis; and a more anterior foramen magnum. While the level of bipedality in <em>Salehanthropus<\/em> <em>tchadenisis<\/em> is debated since there are few fossils and no postcranial evidence, <em>Orrorin tugenensis<\/em> and <em>Ardipithecus<\/em> <em>k<\/em><em>adabba <\/em>show clear indications of some of these bipedal trends. However, some retained ancestral traits, such as an opposable hallux in <em>Ardipithecus<\/em>, indicate some retention in climbing ability.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Derived Adaptations: Early Hominin Dention<\/span><\/h2>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">The Importance of Teeth<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Teeth are abundant in the fossil record, primarily because they are already highly mineralized as they are forming, far more so than even bone. Because of this, teeth preserve readily. And, because they preserve readily, they are well-studied and better understood than many skeletal elements. In the sparse hominin (and primate) fossil record, teeth are, in some cases, all we have.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Teeth also reveal a lot about the individual from whom they came. We can tell what they evolved to eat, to which other species they may be closely related, and even, to some extent, the level of sexual dimorphism, or general variability, within a given species. This is powerful information that can be contained in a single tooth. With a little more observation, the wearing patterns on a tooth can tell us about the diet of the individual in the weeks leading up to its death. Furthermore, the way in which a tooth is formed, and the timing of formation, can reveal information about changes in diet (or even mobility) over infancy and childhood, using isotopic analyses. When it comes to our earliest hominin relatives, this information is vital for understanding how they lived.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The purpose of comparing different hominin species is to better understand the functional morphology as it applies to dentition. In this, we mean that the morphology of the teeth or masticatory system (which includes jaws) can reveal something about the way in which they were used and, therefore, the kinds of foods these hominins ate. When comparing the features of hominin groups, it is worth considering modern analogues (i.e., animals with which to compare) to make more appropriate assumptions about diet. In this way, hominin dentition is often compared with that of chimpanzees and gorillas (our close ape relatives), as well as with that of modern humans.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The most divergent group, however, is humans. Humans around the world have incredibly varied diets. Among hunter-gatherers, it can vary from a honey- and plant-rich diet, as seen in the Hadza in Tanzania, to a diet almost entirely reliant on animal fat and protein, as seen in Inuits in polar regions of the world. We are therefore considered generalists, more general than the largely <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1342\"><strong>frugivorous<\/strong><\/a> (fruit-eating) chimpanzee or the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1344\"><strong>folivorous<\/strong><\/a> (foliage-eating) gorilla, as discussed in Chapter 5.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">One way in which all humans are similar is our reliance on the processing of our food. We cut up and tear meat with tools using our hands, instead of using our front teeth (incisors and canines). We smash and grind up hard seeds, instead of crushing them with our hind teeth (molars). This means that, unlike our ape relatives, we can rely more on developing tools to navigate our complex and varied diets. <span style=\"text-decoration: underline\">(We could say)<\/span> Our brain, therefore, is our primary masticatory organ. Evolutionarily, our teeth have reduced in size and our faces are flatter, or more <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1346\">orthognathic<\/a>, <\/strong>partially in response to our increased reliance on our hands and brain to process food. Similarly, a reduction in teeth and a more generalist dental morphology could also indicate an increase in softer and more variable foods, such as the inclusion of more meat. <span style=\"background-color: #ffff00\">These trends begin early on in our evolution.<\/span> The link has been made between some of the earliest evidence for stone tool manufacture, the earliest members of our genus, and the features that we associate with these specimens.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">General Dental Trends in Early Hominins<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Several trends are visible in the dentition of early hominins. However, all tend to have the same <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1348\"><strong>dental formula<\/strong><\/a>. The dental formula tells us how many of each tooth type are present in each quadrant of the mouth. Going from the front of the mouth, this includes the square, flat <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1160\">incisors<\/a><\/strong>; the pointy <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1166\">canines<\/a><\/strong>; the small, flatter <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1350\"><strong>premolars<\/strong><\/a>; and the larger hind <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1164\">molars<\/a><\/strong>. In many primates, from Old World monkeys to great apes, the typical dental formula is 2:1:2:3. This means that if we divide the mouth into quadrants, each has two incisors, one canine, two premolars, and three molars. The eight teeth per quadrant total 32 teeth in all (although some humans have fewer teeth due to the absence of their wisdom teeth, or third molars).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 380px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image24.png\" alt=\"Anterior view of the lower face of a person showing their teeth.\" width=\"380\" height=\"253\" \/><figcaption class=\"wp-caption-text\">Figure 9.10: In humans, our canines are often a similar size to our incisors. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Adult_human_teeth.jpg\">Adult human teeth<\/a> by <a href=\"https:\/\/www.genusfotografen.se\/\">Genusfotografen<\/a> (Tomas Gunnarsson) through <a href=\"https:\/\/wikimedia.se\/\">Wikimedia Sverige<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p><span style=\"color: #000000\">The morphology of the individual teeth is where we see the most change. Among primates, large incisors are associated with food procurement or preparation (such as biting small fruits), while small incisors indicate a diet that may contain small seeds or leaves (where the preparation is primarily in the back of the mouth). Most hominins have relatively large, flat, vertically aligned incisors that <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1396\">occlude<\/a> <\/strong>(touch) relatively well, forming a \u201cbite.\u201d This differs from, for instance, the orangutan, whose teeth stick out (i.e.<em>,<\/em> are <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1352\"><strong>procumbent<\/strong><\/a>).<\/span><\/p>\n<p>&nbsp;<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">While the teeth are often aligned with diet, the canines may be misleading in that regard. We tend to associate pointy, large canines with the ripping required for meat, and the reduction (or, in some animals, the absence) of canines as indicative of herbivorous diets. In humans, our canines are often a similar size to our incisors and therefore considered <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1398\">incisiform<\/a><\/strong> (Figure 9.10). However, our closest relatives all have very long, pointy canines, particularly on their upper dentition. This is true even for the gorilla, which lives almost exclusively on plants. The canines in these instances reveal more about social structure and sexual dimorphism than diet, as large canines often signal <span style=\"background-color: #ffff00\">dominance<\/span>.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Early on in human evolution, we see a reduction in canine size. <em>Sahelanthropus tchadensis<\/em> and <em>Orrorin tugenensis<\/em> both have smaller canines than those in extant great apes, yet the canines are still larger and pointier than those in humans or more recent hominins. <span style=\"background-color: #ffff00\">This implies strongly that, over evolutionary time, the need for display and dominance among males has reduced, as has our sexual dimorphism.<\/span> In <em>Ardipithecus ramidus<\/em>, there is no obvious difference between male and female canine size, yet they are still slightly larger and pointier than in modern humans. This implies a less sexually dimorphic social structure in the earlier hominins relative to modern-day chimpanzees and gorillas.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Along with a reduction in canine size is the reduction or elimination of a canine <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1172\">diastema<\/a>:<\/strong> a gap between the teeth on the mandible that allows room for elongated teeth on the maxilla to \u201cfit\u201d in the mouth. Absence of a diastema is an excellent indication of a reduction in canine size. In animals with large canines (such as baboons), there is also often a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1354\"><strong>honing P3<\/strong><\/a>, where the first premolar (also known as P3 for evolutionary reasons) is triangular in shape, \u201csharpened\u201d by the extended canine from the upper dentition. This is also seen in some early hominins: <em>Ardipithecus<\/em>, for example, has small canines that are almost the same height as its incisors, although still larger than those in recent hominins.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The hind dentition, such as the bicuspid (two cusped) premolars or the much larger molars, are also highly indicative of a generalist diet in hominins. Among the earliest hominins, the molars are larger than we see in our genus, increasing in size to the back of the mouth and angled in such a way from the much smaller anterior dentition as to give these hominins a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1356\"><strong>parabolic<\/strong><\/a> (V-shaped) dental arch. This differs from our living relatives and some early hominins, such as <em>Sahelanthropus<\/em>, whose molars and premolars are relatively parallel between the left and right sides of the mouth, creating a U-shape.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Among more recent early hominins, the molars are larger than those in the earliest hominins and far larger than those in our own genus, <em>Homo.<\/em> Large, short molars with thick <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1358\"><strong>enamel<\/strong><\/a> allowed our early cousins to grind fibrous, coarse foods, such as sedges, which require plenty of chewing. This is further evidenced in the low <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1176\">cusps<\/a>,<\/strong> or ridges, on the teeth, which are ideal for chewing. In our genus, the hind dentition is far smaller than in these early hominins. Our teeth also have medium-size cusps, which allow for both efficient grinding and tearing\/shearing meats.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Understanding the dental morphology has allowed researchers to extrapolate very specific behaviors of early hominins. It is worth noting that while teeth preserve well and are abundant, a slew of other morphological traits additionally provide evidence for many of these hypotheses. Yet there are some traits that are ambiguous. For instance, while there are definitely high levels of sexual dimorphism in <em>Au. afarensis<\/em>, discussed in the next section, the canine teeth are reduced in size, implying that while canines may be useful indicators for sexual dimorphism, it is also worth considering other evidence.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff99cc\">In summary, trends among early hominins include a reduction in procumbency, reduced hind dentition (molars and premolars), a reduction in canine size (more incisiform with a lack of canine diastema and honing P3), flatter molar cusps, and thicker dental enamel. All early hominins have the ancestral dental formula of 2:1:2:3. These trends are all consistent with a generalist diet, incorporating more fibrous foods.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Special Topic: Contested Species<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Many named species are highly debated and argued to have specimens associated with a more variable <em>Au. afarensis <\/em>or <em>Au. anamensis<\/em> species. Sometimes these specimens are dated to times when, or found in places in which, there are \u201cgaps\u201d in the palaeoanthropological record. These are argued to represent chronospecies or variants of <em>Au. afarensis<\/em>. However, it is possible that, with more discoveries, the distinct species types will hold.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><strong><em>Australopithecus bahrelghazali<\/em><\/strong> is dated to within the time period of <em>Au. afarensi<\/em>s (3.6 mya; Brunet et al. 1995) and was the first Australopithecine to be discovered in Chad in central Africa. Researchers argue that the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1360\">holotype<\/a><\/strong>, whom discoverers have named \u201cAbel,\u201d falls under the range of variation of <em>Au. afarensis<\/em> and therefore that <em>A. bahrelghazali<\/em> does not fall into a new species (Lebatard et al. 2008). If \u201cAbel\u201d is a member of <em>Au. afarensis<\/em>, the geographic range of the species would be greatly extended.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">On a different note, <strong><em>Australopithecus <\/em><\/strong><strong><em>deyiremada<\/em><\/strong> (meaning \u201cclose relative\u201d in the Ethiopian language of Afar) is dated to 3.5 mya to 3.3 mya and is based on fossil mandible bones discovered in 2011 in Woranso-Mille (in the Afar region of Ethiopia) by Yohannes Haile-Selassie, an Ethiopian paleoanthropologist (Haile-Selassie et al. 2019). The discovery indicated, in contrast to <em>Au. afarensis<\/em>, smaller teeth with thicker enamel (potentially suggesting a harder diet) as well as a larger mandible and more projecting cheekbones. This find may be evidence that more than one closely related hominin species occupied the same region at the same temporal period (Haile-Selassie et al. 2015; Spoor 2015) or that other <em>Au. afarensis<\/em> specimens have been incorrectly designated. However, others have argued that this species has been prematurely identified and that more evidence is needed before splitting the taxa, since the variation appears subtle and may be due to slightly different niche occupations between populations over time.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><strong><em>Australopithecus garhi<\/em><\/strong> is another species found in the Middle Awash region of Ethiopia. It is currently dated to 2.5 mya (younger than <em>Au. afarensis<\/em>). Researchers have suggested it fills in a much-needed temporal \u201cgap\u201d between hominin finds in the region, with some anatomical differences, such as a relatively large cranial capacity (450 cc) and larger hind dentition than seen in other gracile Australopithecines. Similarly, the species has been argued to have longer hind limbs than <em>Au. afarensis<\/em>, although it was still able to move arboreally (Asfaw et al. 1999). However, this species is not well documented or understood and is based on only several fossil specimens. More astonishingly, crude stone tools resembling Oldowan (which will be described later) have been found in association with <em>Au. garhi<\/em>. While lacking some of the features of the Oldowan, this is one of the earliest technologies found in direct association with a hominin.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><strong><em>Kenyanthopus<\/em><\/strong><strong><em> platyops<\/em><\/strong> (the name \u201cplatyops\u201d refers to its flatter-faced appearance) is a highly contested genus\/species designation of a specimen (KNM-WT 40000) from Lake Turkana in Kenya, discovered by Maeve Leakey in 1999 (Figure 9.11). Dated to between 3.5 mya and 3.2 mya, some have suggested this specimen is an <em>Australopithecus<\/em>, perhaps even <em>Au.<\/em> <em>afarensis<\/em> (with a brain size which is difficult to determine, yet appears small), while still others have placed this specimen in <em>Homo <\/em>(small dentition and flat-orthognathic face). While taxonomic placing of this species is quite divided, the discoverers have argued that this species is ancestral to <em>Homo<\/em>, in particular to <em>Homo <\/em><em>ruldolfensis<\/em> (Leakey et al. 2001). Some researchers have additionally associated the earliest tool finds from Lomekwi, Kenya, temporally (3.3 mya) and in close geographic proximity to this specimen.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 579px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-292 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.11.jpg\" alt=\"Four views of an ancient skull are shown on a black background.\" width=\"579\" height=\"579\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.11: This specimen, KNM WT 40000 (Kenyanthopus platyops), has small detention, a small brain case, and a relatively flat face. Its genus\/species designation remains contested. Credit: a. <a class=\"rId76\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId77\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 anterior view<\/a> by \u00a9<a class=\"rId78\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId79\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId80\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId81\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId82\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 superior view<\/a> by \u00a9<a class=\"rId83\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId84\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId85\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a class=\"rId86\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId87\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 lateral left view<\/a> by \u00a9<a class=\"rId88\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId89\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId90\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; d. <a class=\"rId91\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId92\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 inferior view<\/a> by \u00a9<a class=\"rId93\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId94\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId95\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">The Genus <em>Australopithecus<\/em><br \/>\n<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The Australopithecines are a diverse group of hominins, comprising various species. <em>Australopithecus<\/em> is the given group or genus name. It stems from the Latin word <em>Australo<\/em>, meaning \u201csouthern,\u201d and the Greek word <em>pithecus,<\/em> meaning \u201cape.\u201d Within this section, we will outline these differing species\u2019 geological and temporal distributions across Africa, unique derived and\/or shared traits, and importance in the fossil record.<\/span><\/p>\n<figure style=\"width: 381px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image37-2.png\" alt=\"A skull has a pronounced sagittal crest, flaring cheekbones, and large hind teeth.\" width=\"381\" height=\"585\" \/><figcaption class=\"wp-caption-text\">Figure 9.12: Robust Australopithecines such as Paranthropus boisei had large molars and chewing muscles. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Paranthropus_boisei_skull.jpg\">Paranthropus boisei skull<\/a> by Durova is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/deed.en\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Between 3 mya and 1 mya, there seems to be differences in dietary strategy between different species of hominins designated as Australopithecines. A pattern of larger posterior dentition (even relative to the incisors and canines in the front of the mouth), thick enamel, and cranial evidence for extremely large chewing muscles is far more pronounced in a group known as the robust australopithecines. This pattern is extreme<span style=\"text-decoration: underline\">ly<\/span> relative to their earlier contemporaries or predecessors, the gracile australopithecines<strong>,<\/strong> and is certainly larger than those seen in early <em>Homo<\/em>, which emerged during this time. This pattern of incredibly large hind dentition (and very small anterior dentition) has led people to refer to robust australopithecines as <strong>megadont<\/strong> hominins (Figure 9.12).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Because of these differences, this section has been divided into \u201cgracile\u201d and \u201crobust\u201d Australopithecines, highlighting the morphological differences between the two groups (which many researchers have designated as separate genera: <em>Australopithecus<\/em> and <em>Paranthropus<\/em>, respectively) and then focusing on the individual species. It is worth noting, however, that not all researchers accept these clades as biologically or genetically distinct, with some researchers insisting that the relative gracile and robust features found in these species are due to parallel evolutionary events toward similar dietary niches.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Despite this genus\u2019 ancestral traits and small cranial capacity, all members show evidence of bipedal locomotion. It is generally accepted that <em>Australopithecus <\/em>species display varying degrees of arborealism along with bipedality.<\/span><\/p>\n<h3 class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Gracile Australopithecines<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">This section describes individual species from across Africa. These species are called \u201c<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1406\">gracile<\/a> <\/strong>australopithecines\u201d because of their smaller and less robust features compared to the divergent \u201c<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1408\"><strong>robust<\/strong><\/a>\u201d group. Numerous Australopithecine species have been named, but some are only based on a handful of fossil finds, whose designations are controversial.<\/span><\/p>\n<h4 class=\"import-Normal\"><em><span style=\"color: #000000\">East African Australopithecines<\/span><\/em><\/h4>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">East African Australopithecines are found throughout the EARS, and they include the earliest species associated with this genus. Numerous fossil-yielding sites, such as Olduvai, Turkana, and Laetoli, have excellent, datable stratigraphy, owing to the layers of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1362\"><strong>volcanic tufts<\/strong> <\/a> that have accumulated over millions of years. <span style=\"background-color: #ff9900\">These tufts may be dated using absolute dating techniques, such as Potassium-Argon dating (described in Chapter 7). This means that it is possible to know a relatively refined date for any fossil if the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1364\"><strong>context<\/strong> <\/a> (i.e., exact location) of that find is known.<\/span> Similarly, comparisons between the faunal assemblages of these stratigraphic layers have allowed researchers to chronologically identify environmental changes.<\/span><\/p>\n<figure style=\"width: 313px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image20-1-1.png\" alt=\"Occlusal view of an Au. anamensis mandible, with relatively large teeth, including canines.\" width=\"313\" height=\"313\" \/><figcaption class=\"wp-caption-text\">Figure 9.13: As seen in this mandible of KNM-KP 29281, Australopithecus anamensis had relatively large canine teeth. Credit: <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20anamensis\/KNM-KP%2029281\">Australopithecus anamensis: KNM-KP 29281 occlusal view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p><span style=\"color: #000000\">The earliest known Australopithecine is dated to 4.2 mya to 3.8 mya. <strong><em>Australopithecus anamensis<\/em><\/strong> (after \u201cAnam,\u201d meaning \u201clake\u201d from the Turkana region in Kenya; Leakey et al. 1995; Patterson and Howells 1967) is currently found from sites in the Turkana region (Kenya) and Middle Awash (Ethiopia; Figure 9.13). Recently, a 2019 find from Ethiopia, named MRD, after Miro Dora where it was found, was discovered by an Ethiopian herder named Ali Bereino. It is one of the most complete cranial finds of this species (Ward et al. 1999). A small brain size (370 cc), relatively large canines, projecting cheekbones, and earholes show more ancestral features as compared to those of more recent Australopithecines. The most important element discovered with this species is a fragment of a tibia (shinbone), which demonstrates features associated with weight transfer during bipedal walking. Similarly, the earliest found hominin femur belongs to this species. Ancestral traits in the upper limb (such as the humerus) indicate some retained arboreal locomotion.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Some researchers suggest that <em>Au. anamensis<\/em> is an intermediate form of the chronospecies that becomes <em>Au. afarensis<\/em>, evolving from <em>Ar. ramidus<\/em>. However, this is debated, with other researchers suggesting morphological similarities and affinities with more recent species instead. Almost 100 specimens, representing over 20 individuals, have been found to date (Leakey et al. 1995; McHenry 2009; Ward et al. 1999).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Au. afarensis<\/em><\/strong> is one of the oldest and most well-known australopithecine species and consists of a large number of fossil remains. <em>Au. afarensis<\/em> (which means \u201cfrom the Afar region\u201d) is dated to between 2.9 mya and 3.9 mya and is found in sites all along the EARS system, in Tanzania, Kenya, and Ethiopia (Figure 9.14). The most famous individual from this species is a partial female skeleton discovered in Hadar (Ethiopia), later nicknamed \u201cLucy,\u201d after the Beatles\u2019 song \u201cLucy in the Sky with Diamonds,\u201d which was played in celebration of the find (Johanson et al. 1978; Kimbel and Delezene 2009). This skeleton was found in 1974 by Donald Johanson and dates to approximately 3.2 mya. In addition, in 2002 a juvenile of the species was found by Zeresenay Alemseged and given the name \u201cSelam\u201d (meaning \u201cpeace,\u201d DIK 1-1), though it is popularly known as \u201cLucy\u2019s Child\u201d or as the \u201cDikika Child\u201d (Alemseged et al. 2006). Similarly, the \u201cLaetoli Footprints\u201d (discussed in Chapter 7; Hay and Leakey 1982; Leakey and Hay 1979) have drawn much attention.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 643px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-295 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.14.jpg\" alt=\"Two images of life-like reconstructions of female and male Au. afarensis.\" width=\"643\" height=\"322\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.14 a-b: Artistic reconstructions of Australopithecus afarensis by artist John Gurche. Female \u201cLucy\u201d is left and a male is on the right. Credit: a. <a class=\"rId106\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Australopithecus afarensis, \u201cLucy,\u201d adult female. Reconstruction based on AL-288-1 by artist John Gurche, front view close-up<\/a> by <a class=\"rId107\" href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?\u201d] is <a class=\"rId108\" href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>; b. <a class=\"rId109\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Australopithecus afarensis, adult male. Reconstruction based on <\/a><a class=\"rId110\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">AL444-2<\/a><a class=\"rId111\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\"> by John Gurche<\/a> by <a class=\"rId112\" href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?\u201d] is <a class=\"rId113\" href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>.<\/figcaption><\/figure>\n<figure style=\"width: 320px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image40.png\" alt=\"A partial skeleton includes bones of the cranium, mandible, and postcranium.\" width=\"320\" height=\"772\" \/><figcaption class=\"wp-caption-text\">Figure 9.15: The humanlike femoral neck, valgus knee, and bowl-shaped hip seen in the \u201cLucy\u201d skeleton indicates that Australopithecus afarensis was bipedal. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Lucy_blackbg.jpg\">Lucy blackbg<\/a> [AL 288-1, Australopithecus afarensis, cast from Museum national d'histoire naturelle, Paris] by 120 is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The canines and molars of <em>Au. afarensis<\/em> are reduced relative to great apes but are larger than those found in modern humans (indicative of a generalist diet); in addition, <em>Au. afarensis <\/em>has a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1366\"><strong>prognathic<\/strong> <\/a> face (the face below the eyes juts anteriorly) and robust facial features that indicate relatively strong chewing musculature (compared with <em>Homo<\/em>) but which are less extreme than in <em>Paranthropus<\/em>. Despite a reduction in canine size in this species, large overall size variation indicates high levels of sexual dimorphism.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Skeletal evidence indicates that this species was bipedal, as its pelvis and lower limb demonstrate a humanlike femoral neck, valgus knee, and bowl-shaped hip (Figure 9.15). <span style=\"background-color: #ccffcc\">More evidence of bipedalism is found in the footprints of this species<\/span>. <em>Au. afarensis<\/em> is associated with the Laetoli Footprints, <span style=\"background-color: #ff9900\">a 24-meter trackway of hominin fossil footprints preserved in volcanic ash discovered by Mary Leakey in Tanzania and dated to 3.5 mya to 3 mya. This set of prints is thought to have been produced by three bipedal individuals as there are no knuckle imprints, no opposable big toes, and a clear arch is present. The infants of this species are thought to have been more arboreal than the adults, as discovered through analyses of the foot bones of the Dikika Child dated to 3.32 mya (Alemseged et al. 2006).<\/span><\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Although not found in direct association with stone tools, potential evidence for cut marks on bones, found at Dikika, and dated to 3.39 mya indicates a possible temporal\/ geographic overlap between meat eating, tool use, and this species. However, this evidence is fiercely debated. Others have associated the cut marks with the earliest tool finds from Lomekwi, Kenya, temporally (3.3 mya) and in close geographic proximity to this species.<\/span><\/p>\n<h4 class=\"import-Normal\"><em><span style=\"color: #000000\">South African Australopithecines<\/span><\/em><\/h4>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Since the discovery of the Taung Child, there have been numerous Australopithecine discoveries from the region known as \u201cThe Cradle of Humankind,\u201d which was recently given UNESCO World Heritage Site status as \u201cThe Fossil Hominid Sites of South Africa.\u201d The limestone caves found in the Cradle allow for the excellent preservation of fossils. Past animals navigating the landscape and falling into cave openings, or caves used as dens by carnivores, led to the accumulation of deposits over millions of years. Many of the hominin fossils, encased in <strong>breccia<\/strong> (hard, calcareous sedimentary rock), are recently exposed from limestone quarries mined in the previous century. This means that extracting fossils requires excellent and detailed exposed work, often by a team of skilled technicians.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">While these sites have historically been difficult to date, with mixed assemblages accumulated over large time periods, advances in techniques such as uranium-series dating have allowed for greater accuracy. Historically, the excellent faunal record from East Africa has been used to compare sites based on <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1368\"><strong>relative dating<\/strong>,<\/a> whereby environmental and faunal changes and extinction events allow us to know which hominin finds are relatively younger or older than others.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The discovery of the Taung Child in 1924 (discussed in the Special Topic box \u201cThe Taung Child\u201d below) shifted the focus of palaeoanthropological research from Europe to Africa, although acceptance of this shift was slow (Broom 1947; Dart 1925). The species to which it is assigned, <strong><em>Australopithecus africanus<\/em><\/strong> (name meaning \u201cSouthern Ape of Africa\u201d), is currently dated to between 3.3 mya and 2.1 mya (Pickering and Kramers 2010), with discoveries from Sterkfontein, Taung, Makapansgat, and Gladysvale in South Africa (Figure 9.16). A relatively large brain (400 cc to 500 cc), small canines without an associated diastema, and more rounded cranium and smaller teeth than <em>Au. afarensis<\/em> indicate some derived traits. Similarly, the postcranial remains (in particular, the pelvis) indicate bipedalism. However, the sloping face and curved phalanges (indicative of retained arboreal locomotor abilities) show some ancestral features. Although not in direct association with stone tools, a 2015 study noted that the trabecular bone morphology of the hand was consistent with forceful tool manufacture and use, suggesting potential early tool abilities.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 570px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image19-1.png\" alt=\"A life-like reconstruction of the face of Australopithecus africanus, smiling in anterior view.\" width=\"570\" height=\"570\" \/><figcaption class=\"wp-caption-text\">Figure 9.16: An artistic reconstruction of Australopithecus africanus by John Gurche. Credit: <a href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Australopithecus africanus. Reconstruction based on STS 5 by John Gurche <\/a>by <a href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?] is <a href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Another famous <em>Au. africanus<\/em> skull (the skull of \u201cMrs. Ples\u201d) was previously attributed to <em>Plesianthropus transvaalensis<\/em><em>, <\/em>meaning \u201cnear human from the Transvaal,\u201d the old name for Gauteng Province, South Africa (Broom 1947, 1950). The name was shortened by contemporary journalists to \u201cPles\u201d (Figure 9.17). Due to the prevailing mores of the time, the assumed female found herself married, at least in name, and has become widely known as \u201cMrs. Ples.\u201d It was later reassigned to <em>Au. africanus<\/em> and is now argued by some to be a young male rather than an adult female cranium (Thackeray 2000, Thackeray et al. 2002).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 548px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-298 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.17.jpg\" alt=\"Four views of an ancient skull are shown on a black background.\" width=\"548\" height=\"548\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.17: The \u201cMrs. Ples\u201d brain case is small in size (like apes) but its face is less prognathic; its foramen magnum is positioned more like a modern human than an African apes. Credit: a. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 anterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 posterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 superior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; and d. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 lateral right view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In 2008, nine-year-old Matthew Berger, son of paleoanthropologist Lee Berger, noted a clavicle bone in some leftover mining breccia in the Malapa Fossil Site (South Africa). After rigorous studies, the species, <strong><em>Australopithecus sediba<\/em><\/strong> (meaning \u201cfountain\u201d or \u201cwellspring\u201d in the South African language of Sesotho), was named in 2010 (Figure 9.18; Berger et al. 2010). The first type specimen belongs to a juvenile male, Karabo (MH1), but the species is known from at least six partial skeletons, from infants through adults. These specimens are currently dated to 1.97 mya (Dirks et al. 2010). The discoverers have argued that <em>Au. sediba<\/em> shows mosaic features between <em>Au. africanus<\/em> and the genus, <em>Homo<\/em>, which potentially indicates a transitional species, although this is heavily debated. These features include a small brain size (<em>Australopithecus<\/em>-like; 420 cc to 450 cc) but gracile mandible and small teeth (<em>Homo<\/em>-like). Similarly, the postcranial skeletons are also said to have mosaic features: scientists have interpreted this mixture of traits (such as a robust ankle but evidence for an arch in the foot) as a transitional phase between a body previously adapted to arborealism (particularly in evidence from the bones of the wrist) to one that adapted to bipedal ground walking. Some researchers have argued that <em>Au. sediba<\/em> shows a modern hand morphology (shorter fingers and a longer thumb), indicating that adaptations to tool manufacture and use may be present in this species.<\/span><\/p>\n<figure style=\"width: 531px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image17-1.png\" alt=\"A beige-colored skull with no mandible on a black background has some missing teeth.\" width=\"531\" height=\"400\" \/><figcaption class=\"wp-caption-text\">Figure 9.18: Australopithecus sediba shows mosaic features between Au. africanus and Homo. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Australopithecus_sediba.JPG\">Australopithecus sediba<\/a>, photo by Brett Eloff courtesy <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Profberger\">Profberger<\/a> and <a href=\"https:\/\/en.wikipedia.org\/wiki\/University_of_the_Witwatersrand\">Wits University<\/a>, is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Another famous Australopithecine find from South Africa is that of the nearly complete skeleton now known as \u201cLittle Foot\u201d (Clarke 1998, 2013). Little Foot (StW 573) is potentially the earliest dated South African hominin fossil, dating to 3.7 mya, based on radiostopic techniques, although some argue that it is younger than 3 mya (Pickering and Kramers 2010). The name is jokingly in contrast to the cryptid species \u201cbigfoot\u201d and is named because the initial discovery of four ankle bones indicated bipedality. Little Foot was discovered by Ron Clarke in 1994, when he came across the ankle bones while sorting through monkey fossils in the University of Witwatersrand collections (Clarke and Tobias 1995). He asked Stephen Motsumi and Nkwane Molefe to identify the known records of the fossils, which allowed them to find the rest of the specimen within just days of searching the Sterkfontein Caves\u2019 Silberberg Grotto.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The discoverers of Little Foot insist that other fossil finds, previously identified as <em>Au. Africanus<\/em>, be placed in this new species based on shared ancestral traits with older East African Australopithecines (Clarke and Kuman 2019). These include features such as a relatively large brain size (408 cc), robust zygomatic arch, and a flatter midface. Furthermore, the discoverers have argued that the heavy anterior dental wear patterns, relatively large anterior dentition, and smaller hind dentition of this specimen more closely resemble that of <em>Au. anamensis<\/em> or <em>Au. afarensis<\/em>. It has thus been placed in the species <strong><em>Australopithecus prometheus<\/em><\/strong>. This species name refers to a previously defunct taxon named by Raymond Dart. The species designation was, through analyzing Little Foot, revived by Ron Clarke, who insists that many other fossil hominin specimens have prematurely been placed into <em>Au. africanus<\/em>. Others say that it is more likely that <em>Au. africanus<\/em> is a more variable species and not representative of two distinct species.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\"><em>Paranthropus<\/em> \u201cRobust\u201d Australopithecines<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In the robust australopithecines, the specialized nature of the teeth and masticatory system, such as flaring zygomatic arches (cheekbones), accommodate very large temporalis (chewing) muscles. These features also include a large, broad, dish-shaped face and and a large mandible with extremely large posterior dentition (referred to as megadonts) and hyper-thick enamel (Kimbel 2015; Lee-Thorp 2011; Wood 2010). Research has revolved around the shared adaptations of these \u201crobust\u201d australopithecines, linking their morphologies to a diet of hard and\/or tough foods (Brain 1967; Rak 1988). Some argued that the diet of the robust australopithecines was so specific that any change in environment would have accelerated their extinction. The generalist nature of the teeth of the gracile australopithecines, and of early <em>Homo<\/em>, would have made them more capable of adapting to environmental change. However, some have suggested that the features of the robust australopithecines might have developed as an effective response to what are known as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1370\"><strong>fallback <\/strong><strong>foods<\/strong><\/a> in hard times rather than indicating a lack of adaptability.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are currently three widely accepted robust australopithecus or, <em>Paranthropus<\/em>, species: <em>P. aethiopic<\/em><em>us<\/em>, which has more ancestral traits, and <em>P. boisei and P. robustus<\/em>, which are more derived in their features (Strait et al. 1997; Wood and Schroer 2017). These three species have been grouped together by a majority of scholars as a single genus as they share more derived features (are more closely related to each other; or, in other words, are <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1372\"><strong>monophyletic<\/strong><\/a>) than the other australopithecines (Grine 1988; Hlazo 2015; Strait et al. 1997; Wood 2010 ). While researchers have mostly agreed to use the umbrella term <em>Paranthropus<\/em>, there are those who disagree (Constantino and Wood 2004, 2007; Wood 2010).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">As a collective, this genus spans 2.7 mya to 1.0 mya, although the dates of the individual species differ. The earliest of the Paranthropus species, <strong><em>Paranthropus aethiopicus<\/em><\/strong>, is dated to between 2.7 mya and 2.3 mya and currently found in Tanzania, Kenya, and Ethiopia in the EARS system (Figure 9.19; Constantino and Wood 2007; Hlazo 2015; Kimbel 2015; Walker et al. 1986; White 1988). It is well known because of one specimen known as the \u201cBlack Skull\u201d (KNM\u2013WT 17000), so called because of the mineral manganese that stained it black during fossilization (Kimbel 2015). As with all robust Australopithecines, <em>P. aethiopicus<\/em> has the shared derived traits of large, flat premolars and molars; large, flaring zygomatic arches for accommodating large chewing muscles (the temporalis muscle); a sagittal crest (ridge on the top of the skull) for increased muscle attachment of the chewing muscles to the skull; and a robust mandible and supraorbital torus (brow ridge). However, only a few teeth have been found. A proximal tibia indicates bipedality and similar body size to <em>Au. afarensis<\/em>. In recent years, researchers have discovered and assigned a proximal tibia and juvenile cranium (L.338y-6) to the species (Wood and Boyle 2016).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 666px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-300 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.19.jpg\" alt=\"Five views of a beige partial skull on a black background.\" width=\"666\" height=\"444\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.19: The \u201cBlack Skull\u201d (Paranthropus aethiopicus) had a large sagittal crest and large, flared zygomatic arches that indicate it had large chewing muscles and a powerful biting force. Credit: a. <a class=\"rId156\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId157\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 anterior view<\/a> by \u00a9<a class=\"rId158\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId159\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId160\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId161\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId162\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 lateral right view<\/a> by \u00a9<a class=\"rId163\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId164\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId165\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a class=\"rId166\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId167\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 superior view<\/a> by \u00a9<a class=\"rId168\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId169\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId170\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; d. <a class=\"rId171\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId172\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 posterior view<\/a> by \u00a9<a class=\"rId173\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId174\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId175\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; e. <a class=\"rId176\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId177\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 inferior view<\/a> by \u00a9<a class=\"rId178\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId179\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId180\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">First attributed as <em>Zinjanthropus boisei<\/em> (with the first discovery going by the nickname \u201cZinj\u201d or sometimes \u201cNutcracker Man\u201d), <strong><em>Paranthropus boisei<\/em><\/strong> was discovered in 1959 by Mary Leakey (see Figure 9.20 and 9.21; Hay 1990; Leakey 1959). This \u201crobust\u201d australopith species is distributed across countries in East Africa at sites such as Kenya (Koobi Fora, West Turkana, and Chesowanja), Malawi (Malema-Chiwondo), Tanzania (Olduvai Gorge and Peninj), and Ethiopia (Omo River Basin and Konso). The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1374\">hypodigm<\/a><\/strong>, sample of fossils whose features define the group, has been found by researchers to date to roughly 2.4 mya to 1.4 mya. Due to the nature of its exaggerated, larger, and more robust features, <em>P. boisei <\/em>has been termed <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1376\"><strong>hyper-robust<\/strong><\/a>\u2014that is, even more heavily built than other robust species, with very large, flat posterior dentition (Kimbel 2015). Tools dated to 2.5 mya in Ethiopia have been argued to possibly belong to this species. Despite the cranial features of <em>P. boisei<\/em> indicating a tough diet of tubers, nuts, and seeds, isotopes indicate a diet high in C4 foods (e.g., grasses, such as sedges). Another famous specimen from this species is the Peninj mandible from Tanzania, found in 1964 by Kimoya Kimeu.<\/span><\/p>\n<figure style=\"width: 557px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image14-1.png\" alt=\"Life-like reconstruction of Paranthropus boisei.\" width=\"557\" height=\"557\" \/><figcaption class=\"wp-caption-text\">Figure 9.20: Artistic reconstruction of a Paranthropus boisei, male, by John Gurche. Credit: <a href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Paranthropus boisei, male. Reconstruction based on OH 5 and KNM-ER 406 by John Gurche<\/a> by <a href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?\u201d] is <a href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>.<\/figcaption><\/figure>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 565px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-302 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.21.jpg\" alt=\"Three views of an ancient skull are shown on a black background.\" width=\"565\" height=\"565\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.21: \u201cNutcracker Man\u201d (Paranthropus boisei) had hyper-robust features including very large dentition, flaring zygomatic arches, a broad concave face. It had a powerful and extremely efficient chewing force. Credit: <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20boisei\/OH%205\">Paranthropus boisei: OH 5 anterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20boisei\/OH%205\">Paranthropus boisei: OH 5 inferior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20boisei\/OH%205\">Paranthropus boisei: OH 5 posterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Paranthropus robustus<\/em><\/strong> was the first taxon to be discovered within the genus in Kromdraai B by a schoolboy named Gert Terblanche; subsequent fossil discoveries were made by researcher Robert Broom in 1938 (Figure 9.22; Broom 1938a, 1938b, 1950), with the holotype specimen TM 1517 (Broom 1938a, 1938b, 1950; Hlazo 2018). <em>Paranthropus robustus<\/em> dates approximately from 2.0 mya to 1 mya and is the only taxon from the genus to be discovered in South Africa. Several of these fossils are fragmentary in nature, distorted, and not well preserved because they have been recovered from quarry breccia using explosives. <em>P. robustus<\/em> features are neither as \u201chyper-robust\u201d as <em>P. boisei<\/em> nor as ancestral as <em>P. aethiopicus<\/em>; instead, they have been described as being less derived, more general features that are shared with both East African species (e.g., the sagittal crest and zygomatic flaring; Rak 1983; Walker and Leakey 1988). Enamel hypoplasia is also common in this species, possibly because of instability in the development of large, thick-enameled dentition.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 572px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-303 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.22.jpg\" alt=\"Four views of a beige-colored skull are shown on a black background.\" width=\"572\" height=\"619\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.22: SK 48, a Paranthropus robustus specimen, had less derived, more general features that were not as robust as P. boisei and not as ancestral as P. aethiopicus. Credit: a. <a class=\"rId208\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId209\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 anterior view<\/a> by \u00a9<a class=\"rId210\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId211\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId212\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId213\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId214\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 superior view<\/a> by \u00a9<a class=\"rId215\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId216\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId217\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a class=\"rId218\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId219\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 inferior view<\/a> by \u00a9<a class=\"rId220\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId221\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId222\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; d. <a class=\"rId223\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId224\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 lateral left view<\/a> by \u00a9<a class=\"rId225\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId226\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId227\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Comparisons between Gracile and Robust Australopiths<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Comparisons between gracile and robust australopithecines may indicate different phylogenetic groupings or parallel evolution in several species. In general, the robust australopithecines have large temporalis (chewing) muscles, as indicated by flaring zygomatic arches, sagittal crests, and robust mandibles (jawbones). Their hind dentition is large (megadont), with low cusps and thick enamel. Within the gracile australopithecines, researchers have debated the relatedness of the species, or even whether these species should be lumped together to represent more variable or polytypic species. Often researchers will attempt to draw chronospecific trajectories, with one taxon said to evolve into another over time.<\/span><\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Special Topic: The Taung Child<\/span><\/h2>\n<figure id=\"attachment_304\" aria-describedby=\"caption-attachment-304\" style=\"width: 570px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-304 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.23.jpg\" alt=\"An ancient skull in anterior and lateral views. One view shows an imprint of the brain.\" width=\"570\" height=\"285\" \/><figcaption id=\"caption-attachment-304\" class=\"wp-caption-text\">Figure 9.23: The Taung Child has a nearly complete face, mandible, and partial endocranial cast. Credit: a. <em>A<\/em><a class=\"rId230\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\"><em>ustralopithecus africanus<\/em><\/a><a class=\"rId231\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\">: Taung 1 anterior view<\/a> by \u00a9<a class=\"rId232\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId233\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId234\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId235\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\"><em>australopithecus africanus<\/em><\/a><a class=\"rId236\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\">: Taung 1 lateral right view<\/a> by \u00a9<a class=\"rId237\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId238\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId239\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">The well-known fossil of a juvenile <em>Australopithecine<\/em>, the \u201cTaung Child,\u201d was the first early hominin evidence ever discovered and was the first to demonstrate our common human heritage in Africa (Figure 9.23; Dart 1925). The tiny facial skeleton and natural endocast were discovered in 1924 by a local quarryman in the North West Province in South Africa and were painstakingly removed from the surrounding cement-like breccia by Raymond Dart using his wife\u2019s knitting needles. When first shared with the scientific community in 1925, it was discounted as being nothing more than a young monkey of some kind. Prevailing biases of the time made it too difficult to contemplate that this small-brained hominin could have anything to do with our own history. The fact that it was discovered in Africa simply served to strengthen this bias.<\/span><\/p>\n<\/div>\n<p>&nbsp;<\/p>\n<h2><span style=\"color: #000000\">Early Tool Use and Technology<br \/>\n<\/span><\/h2>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Early Stone Age Technology (ESA)<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1378\">Early Stone Age (ESA)<\/a><\/strong> marks the beginning of recognizable technology made by our human ancestors. Stone-tool (or <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1380\"><strong>lithic<\/strong><\/a>) technology is defined by the fracturing of rocks and the manufacture of tools through a process called <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1382\"> <strong>knapping<\/strong><\/a>. The Stone Age lasted for more than 3 million years and is broken up into chronological periods called the Early (ESA), Middle (MSA), and Later Stone Ages (LSA). Each period is further broken up into a different <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1384\">techno-complex<\/a><\/strong>, a term encompassing multiple <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1386\">assemblages<\/a><\/strong> (collections of artifacts) that share similar traits in terms of artifact production and morphology. The ESA spanned the largest technological time period of human innovation from over 3 million years ago to around 300,000 years ago and is associated almost entirely with hominin species prior to modern <em>Homo sapiens. <\/em>As the ESA advanced, stone tool makers (known as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1390\"><strong>knappers<\/strong><\/a>) began to change the ways they detached <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1388\">flakes<\/a><\/strong> and eventually were able to shape artifacts into functional tools. These advances in technology go together with the developments in human evolution and cognition, dispersal of populations across the African continent and the world, and climatic changes.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In order to understand the ESA, it is important to consider that not all assemblages are exactly the same within each techno-complex: one can have multiple phases and traditions at different sites (Lombard et al. 2012). However, there is an overarching commonality between them. Within stone tool assemblages, both flakes or <strong>cores<\/strong> (the rocks from which flakes are removed) are used as tools.<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1392\"> <strong>Large Cutting Tools (LCTs)<\/strong><\/a> are tools that are shaped to have functional edges. It is important to note that the information presented here is a small fraction of what is known about the ESA, and there are ongoing debates and discoveries within archaeology.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Currently, the oldest-known stone tools, which form the techno-complex the Lomekwian, date to 3.3 mya (Harmand et al. 2015; Toth 1985). They were found at a site called Lomekwi 3 in Kenya. This techno-complex is the most recently defined and pushed back the oldest-known date for lithic technology. There is only one known site thus far and, due to the age of the site, it is associated with species prior to <em>Homo<\/em>, such as <em>Kenyanthropus platyops.<\/em> Flakes were produced through indirect percussion, whereby the knappers held a rock and hit it against another rock resting on the ground. The pieces are very chunky and do not display the same fracture patterns seen in later techno-complexes. Lomekwian knappers likely aimed to get a sharp-edged piece on a flake, which would have been functional, although the specific function is currently unknown.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Stone tool use, however, is not only understood through the direct discovery of the tools. Cut marks on fossilized animal bones may illuminate the functionality of stone tools. In one controversial study in 2010, researchers argued that cut marks on a pair of animal bones from Dikika (Ethiopia), dated to 3.4 mya, were from stone tools. The discoverers suggested that they be more securely associated, temporally, with <em>Au. afarensis<\/em>. However, others have noted that these marks are consistent with teeth marks from crocodiles and other carnivores.<\/span><\/p>\n<figure style=\"width: 324px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image29-1.png\" alt=\"A technical line drawing of an Oldowan chopper.\" width=\"324\" height=\"275\" \/><figcaption class=\"wp-caption-text\">Figure 9.24: Some scholars believe that some genera explored in this chapter were capable of producing more complex stone tools (Oldowan). Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Olduwan_Industry_Chopper_2.jpeg\">Olduwan Industry Chopper 2<\/a> by Emmyanne29 is under a <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/legalcode\">CC0 1.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The Oldowan techno-complex is far more established in the scientific literature (Leakey 1971). It is called the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1394\">Oldowan<\/a><\/strong> because it was originally discovered in Olduvai Gorge, Tanzania, but the oldest assemblage is from Gona in Ethiopia, dated to 2.6 mya (Semaw 2000). The techno-complex is defined as a core and flake industry. Like the Lomekwian, there was an aim to get sharp-edged flakes, but this was achieved through a different production method. Knappers were able to actively hold or manipulate the core being knapped, which they could directly hit using a hammerstone. This technique is known as free-hand percussion, and it demonstrates an understanding of fracture mechanics. It has long been argued that the Oldowan hominins were skillful in tool manufacture.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Because Oldowan knapping requires skill, earlier researchers have attributed these tools to members of our genus, <em>Homo<\/em>. However, some have argued that these tools are in more direct association with hominins in the genera described in this chapter (Figure 9.24).<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Invisible Tool Manufacture and Use<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The vast majority of our understanding of these early hominins comes from fossils and reconstructed paleoenvironments. It is only from 3 mya when we can start \u201clooking into their minds\u201d and lifestyles by analyzing their manufacture and use of stone tools. However, the vast majority of tool use in primates (and, one can argue, in humans) is not with durable materials like stone. All of our extant great ape relatives have been observed using sticks, leaves, and other materials for some secondary purpose (to wade across rivers, to \u201cfish\u201d for termites, or to absorb water for drinking). It is possible that the majority of early hominin tool use and manufacture may be invisible to us because of this preservation bias.<\/span><\/p>\n<h2 class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000;background-color: #ff00ff\">Chapter Summary<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000;background-color: #ff00ff\">The fossil record of our earliest hominin relatives has allowed paleoanthropologists to unpack some of the mysteries of our evolution. We now know that traits associated with bipedalism evolved before other \u201chuman-like\u201d traits, even though the first hominins were still very capable of arboreal locomotion. We also know that, for much of this time, hominin taxa were diverse in the way they looked and what they ate, and they were widely distributed across the African continent. And we know that the environments in which these hominins lived underwent many changes over this time during several warming and cooling phases.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;background-color: #ff00ff\">Yet this knowledge has opened up many new mysteries. We still need to better differentiate some taxa. In addition, there are ongoing debates about why certain traits evolved and what they meant for the extinction of some of our relatives (like the robust australopiths). The capabilities of these early hominins with respect to tool use and manufacture is also still uncertain.<\/span><\/p>\n<h2 class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Hominin Species Summaries<br \/>\n<\/span><\/h2>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Sahelanthropus tchadensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">7 mya to 6 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Chad<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The initial discovery, made in 2001.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">360 cc average<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Smaller than in extant great apes; larger and pointier than in humans. Canines worn at the tips.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A short cranial base and a foramen magnum (hole in which the spinal cord enters the cranium) that is more humanlike in positioning; has been argued to indicate upright walking.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Currently little published postcranial material.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The extent to which this hominin was bipedal is currently heavily debated. If so, it would indicate an arboreal bipedal ancestor of hominins, not a knuckle-walker like chimpanzees.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Orrorin tugenensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">6 mya to 5.7 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Tugen Hills (Kenya)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Original discovery in 2000.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Smaller cheek teeth (molars and premolars) than even more recent hominins (i.e., derived), thick enamel, and reduced, but apelike, canines.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Not many found<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Fragmentary leg, arm, and finger bones have been found. Indicates bipedal locomotion.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Potential toolmaking capability based on hand morphology, but nothing found directly.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">This is the earliest species that clearly indicates adaptations for bipedal locomotion.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Ardipithecus kadabba<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">5.2 mya to 5.8 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Middle Awash (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Discovered by Yohannes Haile-Selassie in 1997.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Larger hind dentition than in modern chimpanzees. Thick enamel and larger canines than in later hominins.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A large hallux (big toe) bone indicates a bipedal \u201cpush off.\u201d<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Faunal evidence indicates a mixed grassland\/woodland environment.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Ardipithecus ramidus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">4.4 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Middle Awash region and Gona (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">A partial female skeleton nicknamed \u201cArdi\u201d (ARA-VP-6\/500) (found in 1994).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">300 cc to 350 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Little differences between the canines of males and females (small sexual dimorphism).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Midfacial projection, slightly prognathic. Cheekbones less flared and robust than in later hominins.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Ardi demonstrates a mosaic of ancestral and derived characteristics in the postcrania. For instance, an opposable big toe similar to chimpanzees (i.e., more ancestral), which could have aided in climbing trees effectively. However, the pelvis and hip show that she could walk upright (i.e., it is derived), supporting her hominin status.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">None directly associated<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">Over 110 specimens from Aramis<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus anamensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">4.2 mya to 3.8 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Turkana region (Kenya); Middle Awash (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A 2019 find from Ethiopia, named MRD.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">370 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Relatively large canines compared with more recent Australopithecines.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Projecting cheekbones and ancestral earholes.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lower limb bones (tibia and femur) indicate bipedality; arboreal features in upper limb bones (humerus) found.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Almost 100 specimens, representing over 20 individuals, have been found to date.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus afarensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">3.9 mya to 2.9 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Afar Region, Omo, Maka, Fejej, and Belohdelie (Ethiopia); Laetoli (Tanzania); Koobi Fora (Kenya)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lucy (discovery: 1974), Selam (Dikika Child, discovery: 2000), Laetoli Footprints (discovery: 1976).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">380 cc to 430 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Reduced canines and molars relative to great apes but larger than in modern humans.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Prognathic face, facial features indicate relatively strong chewing musculature (compared with <em>Homo<\/em>) but less extreme than in <em>Paranthropus<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clear evidence for bipedalism from lower limb postcranial bones. Laetoli Footprints indicate humanlike walking. Dikika Child bones indicate retained ancestral arboreal traits in the postcrania.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">None directly, but close in age and proximity to controversial cut marks at Dikika and early tools in Lomekwi.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Au. afarensis<\/em> is one of the oldest and most well-known australopithecine species and consists of a large number of fossil remains.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus bahrelghazali<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">3.6 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Chad<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">\u201cAbel,\u201d the holotype (discovery: 1995).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Arguably within range of variation of <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus prometheus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">3.7 mya (debated)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">Sterkfontein (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">\u201cLittle Foot\u201d (StW 573) (discovery: 1994)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">408 cc (Little Foot estimate)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">Heavy anterior dental wear patterns, relatively large anterior dentition and smaller hind dentition, similar to <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">Relatively larger brain size, robust zygomatic arch, and a flatter midface.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">The initial discovery of four ankle bones indicated bipedality.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">Highly debated new species designation.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus <\/em><em>deyiremada<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">3.5 mya to 3.3 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Woranso-Mille (Afar region, Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">First fossil mandible bones were discovered in 2011 in the Afar region of Ethiopia by Yohannes Haile-Selassie.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Smaller teeth with thicker enamel than seen in <em>Au. afarensis<\/em>, with a potentially hardier diet.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Larger mandible and more projecting cheekbones than in <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Contested species designation; arguably a member of <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Kenyanthopus<\/em><em> platyops<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">3.5 mya to 3.2 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Lake Turkana (Kenya)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">KNM\u2013WT 40000 (discovered 1999)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Difficult to determine but appears within the range of <em>Australopithecus afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Small molars\/dentition (<em>Homo<\/em>-like characteristic)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Flatter (i.e., orthognathic) face<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Some have associated the earliest tool finds from Lomekwi, Kenya, temporally (3.3 mya) and in close geographic proximity to this species\/specimen.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Taxonomic placing of this species is quite divided. The discoverers have argued that this species is ancestral to <em>Homo<\/em>, in particular to <em>Homo <\/em><em>ruldolfensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Australopithecus africanus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">3.3 mya to 2.1 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Sterkfontein, Taung, Makapansgat, Gladysvale (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Taung Child (discovery in 1994), \u201cMrs. Ples\u201d (discover in 1947), Little Foot (arguable; discovery in 1994).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">400 cc to 500 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Smaller teeth (derived) relative to <em>Au. afarensis<\/em>. Small canines with no diastema.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">A rounder skull compared with <em>Au. afarensis<\/em> in East Africa. A sloping face (ancestral).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Similar postcranial evidence for bipedal locomotion (derived pelvis) with retained arboreal locomotion, e.g., curved phalanges (fingers), as seen in <em>Au. afarensis.<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">None with direct evidence.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">A 2015 study noted that the trabecular bone morphology of the hand was consistent with forceful tool manufacture and use, suggesting potential early tool abilities.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus garhi<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.5 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Middle Awash (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">450 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Larger hind dentition than seen in other gracile Australopithecines.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A femur of a fragmentary partial skeleton, argued to belong to <em>Au. garhi<\/em>, indicates this species may be longer-limbed than <em>Au. afarensis<\/em>, although still able to move arboreally.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Crude stone tools resembling Oldowan (described later) have been found in association with <em>Au. garhi<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">This species is not well documented or understood and is based on only a few fossil specimens.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Paranthropus aethiopicus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.7 mya to 2.3 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">West Turkana (Kenya); Laetoli (Tanzania); Omo River Basin (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The \u201cBlack Skull\u201d (KNM\u2013WT 17000) (discovery 1985).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain Size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">410 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>P. aethiopicus<\/em> has the shared derived traits of large flat premolars and molars, although few teeth have been found.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Large flaring zygomatic arches for accommodating large chewing muscles (the temporalis muscle), a sagittal crest for increased muscle attachment of the chewing muscles to the skull, and a robust mandible and supraorbital torus (brow ridge).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A proximal tibia indicates bipedality and similar size to <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The \u201cBlack Skull\u201d is so called because of the mineral manganese that stained it black during fossilization.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Paranthropus boisei<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.4 mya to 1.4 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Koobi Fora, West Turkana, and Chesowanja (Kenya); Malema-Chiwondo (Malawi), Olduvai Gorge and Peninj (Tanzania); and Omo River basin and Konso (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">\u201cZinj,\u201d or sometimes \u201cNutcracker Man\u201d (OH5), in 1959 by Mary Leakey. The Peninj mandible from Tanzania, found in 1964 by Kimoya Kimeu.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">500 cc to 550 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Very large, flat posterior dentition (largest of all hominins currently known). Much smaller anterior dentition. Very thick dental enamel.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Indications of very large chewing muscles (e.g., flaring zygomatic arches and a large sagittal crest).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Evidence for high variability and sexual dimorphism, with estimates of males at 1.37 meters tall and females at 1.24 meters.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Richard Leakey and Bernard Wood have both suggested that<em> P. boisei<\/em> could have made and used stone tools. Tools dated to 2.5 mya in Ethiopia have been argued to possibly belong to this species.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Despite the cranial features of <em>P. boisei<\/em> indicating a tough diet of tubers, nuts, and seeds, isotopes indicate a diet high in C4 foods (e.g., grasses, such as sedges). This differs from what is seen in<em> P. robustus<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Australopithecus sediba<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">1.97 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Malapa Fossil Site (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Karabo (MH1) (discovery in 2008)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">420 cc to 450 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Small dentition with Australopithecine cusp-spacing.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Small brain size (<em>Australopithecus<\/em>-like) but gracile mandible (<em>Homo<\/em>-like).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Scientists have interpreted this mixture of traits (such as a robust ankle but evidence for an arch in the foot) as a transitional phase between a body previously adapted to arborealism (tree climbing, particularly in evidence from the bones of the wrist) to one that adapted to bipedal ground walking.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">None of direct association, but some have argued that a modern hand morphology (shorter fingers and a longer thumb) means that adaptations to tool manufacture and use may be present in this species.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">It was first discovered through a clavicle bone in 2008 by nine-year-old Matthew Berger, son of paleoanthropologist Lee Berger.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Paranthropus robustus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.3 mya to 1 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kromdraai B, Swartkrans, Gondolin, Drimolen, and Coopers Cave (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">SK48 (original skull)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">410 cc to 530 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Large posterior teeth with thick enamel, consistent with other Robust Australopithecines. Enamel hypoplasia is also common in this species, possibly because of instability in the development of large, thick enameled dentition.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>P. robustus<\/em> features are neither as \u201chyper-robust\u201d as <em>P. boisei<\/em> or as ancestral in features as <em>P. aethiopicus<\/em>. They have been described as less derived, more general features that are shared with both East African species (e.g., the sagittal crest and zygomatic flaring).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Reconstructions indicate sexual dimorphism.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Several of these fossils are fragmentary in nature, distorted, and not well preserved, because they have been recovered from quarry breccia using explosives.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\"><strong><span style=\"color: #000000\">Review Questions<br \/>\n<\/span><\/strong><\/h2>\n<ul>\n<li class=\"import-Normal\"><span style=\"color: #000000\">What is the difference between a \u201cderived\u201d versus an \u201cancestral\u201d trait? Give an example of both, seen in <em>Au. afarensis<\/em>.<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">Which of the paleoenvironment hypotheses have been used to describe early hominin diversity, and which have been used to describe bipedalism?<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">Which anatomical features for bipedalism do we see in early hominins?<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">Describe the dentition of gracile and robust australopithecines. What might these tell us about their diets?<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">List the hominin species argued to be associated with stone tool technologies. Are you convinced of these associations? Why\/why not?<\/span><\/li>\n<\/ul>\n<\/div>\n<h2><span style=\"color: #000000\">Key Terms<\/span><\/h2>\n<p><span style=\"color: #000000\"><strong>Arboreal:<\/strong> Related to trees or woodland.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Aridification:<\/strong> Becoming increasingly arid or dry, as related to the climate or environment.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Aridity Hypothesis:<\/strong> The hypothesis that long-term aridification and expansion of savannah biomes were drivers in diversification in early hominin evolution.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Assemblage:<\/strong> A collection demonstrating a pattern. Often pertaining to a site or region.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Bipedalism:<\/strong> The locomotor ability to walk on two legs.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Breccia:<\/strong> Hard, calcareous sedimentary rock.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Canines:<\/strong> The pointy teeth just next to the incisors, in the front of the mouth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cheek teeth:<\/strong> Or hind dentition (molars and premolars).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Chronospecies:<\/strong> Species that are said to evolve into another species, in a linear fashion, over time.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Clade:<\/strong> A group of species or taxa with a shared common ancestor.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cladistics:<\/strong> The field of grouping organisms into those with shared ancestry.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Context:<\/strong> As pertaining to palaeoanthropology, this term refers to the place where an artifact or fossil is found.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cores:<\/strong> The remains of a rock that has been flaked or knapped.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cusps:<\/strong> The ridges or \u201cbumps\u201d on the teeth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Dental formula:<\/strong> A technique to describe the number of incisors, canines, premolars, and molars in each quadrant of the mouth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Derived traits:<\/strong> Newly evolved traits that differ from those seen in the ancestor.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Diastema:<\/strong> A tooth gap between the incisors and canines.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Early Stone Age (ESA):<\/strong> The earliest-described archaeological period in which we start seeing stone-tool technology.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>East African Rift System (EARS):<\/strong> This term is often used to refer to the Rift Valley, expanding from Malawi to Ethiopia. This active geological structure is responsible for much of the visibility of the paleoanthropological record in East Africa.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Enamel:<\/strong> The highly mineralized outer layer of the tooth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Encephalization:<\/strong> Expansion of the brain.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Extant:<\/strong> Currently living\u2014i.e., not extinct.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Fallback foods:<\/strong> Foods that may not be preferred by an animal (e.g., foods that are not nutritionally dense) but that are essential for survival in times of stress or scarcity.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Fauna:<\/strong> The animals of a particular region, habitat, or geological period.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Faunal assemblages:<\/strong> Collections of fossils of the animals found at a site.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Faunal turnover:<\/strong> The rate at which species go extinct and are replaced with new species.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Flake:<\/strong> The piece knocked off of a stone core during the manufacture of a tool, which may be used as a stone tool.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Flora:<\/strong> The plants of a particular region, habitat, or geological period.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Folivorous:<\/strong> Foliage-eating.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Foramen magnum:<\/strong> The large hole (foramen) at the base of the cranium, through which the spinal cord enters the skull.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Fossil:<\/strong> The remains or impression of an organism from the past.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Frugivorous:<\/strong> Fruit-eating.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Generalist:<\/strong> A species that can thrive in a wide variety of habitats and can have a varied diet.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Glacial:<\/strong> Colder, drier periods during an ice age when there is more ice trapped at the poles.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Gracile:<\/strong> Slender, less rugged, or pronounced features.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hallux:<\/strong> The big toe.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Holotype:<\/strong> A single specimen from which a species or taxon is described or named.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hominin:<\/strong> A primate category that includes humans and our fossil relatives since our divergence from extant great apes.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Honing P3:<\/strong> The mandibular premolar alongside the canine (in primates, the P3), which is angled to give space for (and sharpen) the upper canines.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hyper-robust:<\/strong> Even more robust than considered normal in the Paranthropus genus.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hypodigm:<\/strong> A sample (here, fossil) from which researchers extrapolate features of a population.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Incisiform:<\/strong> An adjective referring to a canine that appears more incisor-like in morphology.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Incisors:<\/strong> The teeth in the front of the mouth, used to bite off food.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Interglacial:<\/strong> A period of milder climate in between two glacial periods.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Isotopes:<\/strong> Two or more forms of the same element that contain equal numbers of protons but different numbers of neutrons, giving them the same chemical properties but different atomic masses.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Knappers:<\/strong> The people who fractured rocks in order to manufacture tools.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Knapping:<\/strong> The fracturing of rocks for the manufacture of tools.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Large Cutting Tool (LCT):<\/strong> A tool that is shaped to have functional edges.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Last Common Ancestor (LCA):<\/strong> The hypothetical final ancestor (or ancestral population) of two or more taxa before their divergence.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Lithic:<\/strong> Relating to stone (here to stone tools).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Lumbar lordosis:<\/strong> The inward curving of the lower (lumbar) parts of the spine. The lower curve in the human S-shaped spine.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Lumpers:<\/strong> Researchers who prefer to lump variable specimens into a single species or taxon and who feel high levels of variation is biologically real.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Megadont:<\/strong> An organism with extremely large dentition compared with body size.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Metacarpals:<\/strong> The long bones of the hand that connect to the phalanges (finger bones).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Molars:<\/strong> The largest, most posterior of the hind dentition.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Monophyletic:<\/strong> A taxon or group of taxa descended from a common ancestor that is not shared with another taxon or group.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Morphology:<\/strong> The study of the form or size and shape of things; in this case, skeletal parts.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Mosaic evolution:<\/strong> The concept that evolutionary change does not occur homogeneously throughout the body in organisms.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Obligate bipedalism:<\/strong> Where the primary form of locomotion for an organism is bipedal.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Occlude:<\/strong> When the teeth from the maxilla come into contact with the teeth in the mandible.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Oldowan:<\/strong> Lower Paleolithic, the earliest stone tool culture.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Orthognathic:<\/strong> The face below the eyes is relatively flat and does not jut out anteriorly.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Paleoanthropologists:<\/strong> Researchers that study human evolution.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Paleoenvironment:<\/strong> An environment from a period in the Earth\u2019s geological past.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Parabolic:<\/strong> Like a parabola (parabola-shaped).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Phalanges:<\/strong> Long bones in the hand and fingers.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Phylogenetics:<\/strong> The study of phylogeny.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Phylogeny:<\/strong> The study of the evolutionary relationships between groups of organisms.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Pliocene:<\/strong> A geological epoch between the Miocene and Pleistocene.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Polytypic:<\/strong> In reference to taxonomy, having two or more group variants capable of interacting and breeding biologically but having morphological population differences.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Postcranium:<\/strong> The skeleton below the cranium (head).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Premolars:<\/strong> The smallest of the hind teeth, behind the canines.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Procumbent:<\/strong> In reference to incisors, tilting forward.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Prognathic:<\/strong> In reference to the face, the area below the eyes juts anteriorly.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Quaternary Ice Age:<\/strong> The most recent geological time period, which includes the Pleistocene and Holocene Epochs and which is defined by the cyclicity of increasing and decreasing ice sheets at the poles.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Relative dating:<\/strong> Dating techniques that refer to a temporal sequence (i.e., older or younger than others in the reference) and do not estimate actual or absolute dates.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Robust:<\/strong> Rugged or exaggerated features.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Site:<\/strong> A place in which evidence of past societies\/species\/activities may be observed through archaeological or paleontological practice.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Specialist:<\/strong> A specialist species can thrive only in a narrow range of environmental conditions or has a limited diet.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Splitters:<\/strong> Researchers who prefer to split a highly variable taxon into multiple groups or species.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Taxa:<\/strong> Plural of taxon, a taxonomic group such as species, genus, or family.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Taxonomy:<\/strong> The science of grouping and classifying organisms.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Techno-complex:<\/strong> A term encompassing multiple assemblages that share similar traits in terms of artifact production and morphology.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Thermoregulation:<\/strong> Maintaining body temperature through physiologically cooling or warming the body.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Ungulates:<\/strong> Hoofed mammals\u2014e.g., cows and kudu.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Volcanic tufts:<\/strong> Rock made from ash from volcanic eruptions in the past.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Valgus knee:<\/strong> The angle of the knee between the femur and tibia, which allows for weight distribution to be angled closer to the point above the center of gravity (i.e., between the feet) in bipeds.<\/span><\/p>\n<h2 class=\"import-Normal\"><strong><span style=\"color: #000000\">About the Authors<br \/>\n<\/span><\/strong><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image31.jpg\" alt=\"A woman with short blonde hair smiles at the camera.\" width=\"311\" height=\"311\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Kerryn Warren, Ph.D.<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Grad Coach International, <a class=\"rId245\" style=\"color: #000000\" href=\"mailto:kerryn.warren@gmail.com\">kerryn.warren@gmail.com<\/a><\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kerryn Warren is a dissertation coach at Grad Coach International and is passionate about stimulating research thinking in students of all levels. She has lectured on multiple topics, including archaeology and human evolution, with her research and science communication interests including hybridization in the hominin fossil record (stemming from research from her Ph.D.) and understanding how evolution is taught in South African schools. She also worked as one of the \u201cUnderground Astronauts,\u201d selected to excavate <em>Homo naledi <\/em>remains from the Rising Star Cave System in the Cradle of Humankind.<\/span><\/p>\n<p>&nbsp;<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image11-2.jpg\" alt=\"A woman with short brown hair smiles at the camera.\" width=\"312\" height=\"306\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">K. Lindsay Hunter, M.A., Ph.D. candidate<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">CARTA, k.lindsay.hunter@gmail.com<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lindsay Hunter is a trained palaeoanthropologist who uses her more than 15 years of experience to make sense of the distant past of our species to build a better future. She received her master\u2019s degree in biological anthropology from the University of Iowa and is completing her Ph.D. in archaeology at the University of the Witwatersrand in Johannesburg, South Africa. She has studied fossil and human bone collections across five continents with major grant support from the National Science Foundation (United States) and the Wenner-Gren Foundation for Anthropological Research. As a National Geographic Explorer, Lindsay developed and managed the National Geographic\u2013sponsored Umsuka Public Palaeoanthropology Project in the Cradle of Humankind World Heritage Site (CoH WHS) in South Africa from within Westbury Township, Johannesburg, between 2016\u20132019. She currently serves as the Community Engagement &amp; Advancement Director for CARTA: The UC San Diego\/Salk Institute Center for Academic Research and Training in Anthropogeny in La Jolla, California.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image25-2.jpg\" alt=\"A woman with black hair stands in a hole in the ground.\" width=\"295\" height=\"339\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Navashni Naidoo, M.Sc.<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">University of Cape Town, nnaidoo2@illinois.edu<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Navashni Naidoo is a researcher at Nelson Mandela University, lecturing on physical geology. She completed her Master\u2019s in Science in Archaeology in 2017 at the University of Cape Town. Her research interests include developing paleoenvironmental proxies suited to the African continent, behavioral ecology, and engaging with community-driven archaeological projects. She has excavated at Stone Age sites across Southern Africa and East Africa. Navashni is currently pursuing a PhD in the Department of Anthropology at the University of Illinois.<\/span><\/p>\n<p>&nbsp;<\/p>\n<p>&nbsp;<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image47.jpg\" alt=\"A man with black hair and dark brown eyes looks at the camera. \" width=\"294\" height=\"294\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Silindokuhle Mavuso, M.Sc.<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">University of Witwatersrand, S.muvaso@ru.ac.za<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Silindokuhle has always been curious about the world around him and how it has been shaped. He is a lecturer at Rhodes University of Witwatersrand (Wits), and conducts research on palaeoenvironmental reconstruction and change of the northeastern Turkana Basin\u2019s Pleistocene sequence. Silindokuhle began his education with a B.Sc. (Geology, Archaeology, and Environmental and Geographical Sciences) from the University of Cape Town before moving to Wits for a B.Sc. Honors (geology and paleontology) and M.Sc. in geology. He is currently concluding his PhD Studies. During this time, he has gained more training as a Koobi Fora Fieldschool fellow (Kenya) as well as an Erasmus Mundus scholar (France). Silindokuhle is a Plio-Pleistocene geologist with a specific focus on identifying and explaining past environments that are associated with early human life and development through time. He is interested in a wide range of disciplines such as micromorphology, sedimentology, geochemistry, geochronology, and sequence stratigraphy. He has worked with teams from significant eastern and southern African hominid sites including Elandsfontein, Rising Star, Sterkfontein, Gondolin, Laetoli, Olduvai, and Koobi Fora.<\/span><\/p>\n<h2 class=\"import-Normal\"><strong><span style=\"color: #000000\">For Further Exploration<br \/>\n<\/span><\/strong><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/humanorigins.si.edu\/evidence\">The Smithsonian Institution website<\/a> hosts descriptions of fossil species, an interactive timeline, and much more.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.maropeng.co.za\/content\/page\/human-evolution\">The Maropeng Museum website<\/a> hosts a wealth of information regarding South African Fossil Bearing sites in the Cradle of Humankind<strong>.<\/strong><\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/perot-museum.imgix.net\/2019-08-naledi-sediba-quick-comparison.pdf\">This quick comparison between <em>Homo naledi<\/em> and <em>Australopithecus sediba<\/em><\/a> from the Perot Museum.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.dropbox.com\/s\/l1d2hv42psj21y9\/Braided%20Stream-1920.mp4?dl=0\">This explanation of the braided stream<\/a> by the Perot Museum.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.hetmp.com\/\">A collation of 3-D files for visualizing<\/a> (or even 3-D printing) for homes, schools, and universities.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.pbslearningmedia.org\/resource\/tdc02.sci.life.evo.lp_humanevo\/human-evolution.\">PBS learning materials<\/a>, including videos and diagrams of the Laetoli footprints, bipedalism, and fossils.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A wealth of <a href=\"https:\/\/australianmuseum.net.au\/learn\/science\/human-evolution\/\">information from the Australian Museum website<\/a>, including species descriptions, family trees, and explanations of bipedalism and diet<strong>.<\/strong><\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\"><strong>References<\/strong><\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Alemseged, Zeresenay, Fred Spoor, William H. Kimbel, Ren\u00e9 Bobe, Denis Geraads, Denn\u00e9 Reed, and Jonathan G. Wynn. 2006. \u201cA Juvenile Early Hominin Skeleton from Dikika, Ethiopia.\u201d <em>Nature<\/em> 443 (7109): 296\u2013301.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Asfaw, Berhane, Tim White, Owen Lovejoy, Bruce Latimer, Scott Simpson, and Gen Suwa. 1999. \u201c<em>Australopithecus garhi<\/em>: A New Species of Early Hominid from Ethiopia.\u201d <em>Science<\/em> 284 (5414): 629\u2013635.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Behrensmeyer, Anna K., Nancy E. Todd, Richard Potts, and Geraldine E. McBrinn. 1997. \u201cLate Pliocene Faunal Turnover in the Turkana Basin, Kenya, and Ethiopia.\u201d <em>Science<\/em> 278 (5343): 637\u2013640.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Berger, Lee R., Darryl J. De Ruiter, Steven E. Churchill, Peter Schmid, Kristian J. Carlson, Paul HGM Dirks, and Job M. Kibii. 2010. \u201c<em>Australopithecus sediba<\/em>: A New Species of <em>Homo<\/em>-like Australopith from South Africa.\u201d <em>Science<\/em> 328 (5975): 195\u2013204.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Bobe, Ren\u00e9, and Anna K. Behrensmeyer. 2004. \u201cThe Expansion of Grassland Ecosystems in Africa in Relation to Mammalian Evolution and the Origin of the Genus <em>Homo<\/em>.\u201d <em>Palaeogeography, Palaeoclimatology, Palaeoecology<\/em> 207 (3\u20134): 399\u2013420.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Brain, C. K. 1967. \u201cThe Transvaal Museum's Fossil Project at Swartkrans.\u201d <em>South African Journal of Science<\/em> 63 (9): 378\u2013384.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1938a. \u201cMore Discoveries of Australopithecus.\u201d <em>Nature<\/em> 141 (1): 828\u2013829.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1938b. \u201cThe Pleistocene Anthropoid Apes of South Africa.\u201d <em>Nature<\/em> 142 (3591): 377\u2013379.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1947. \u201cDiscovery of a New Skull of the South African Ape-Man, Plesianthropus.\u201d <em>Nature<\/em> 159 (4046): 672.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1950. \u201cThe Genera and Species of the South African Fossil Ape-Man.\u201d <em>American Journal of Physical Anthropology<\/em> 8 (1): 1\u201314.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Brunet, Michel, Alain Beauvilain, Yves Coppens, Emile Heintz, Aladji HE Moutaye, and David Pilbeam. 1995. \u201cThe First Australopithecine 2,500 Kilometers West of the Rift Valley (Chad).\u201d <em>Nature<\/em> 378 (6554): 275\u2013273.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Cerling, Thure E., Jonathan G. Wynn, Samuel A. Andanje, Michael I. Bird, David Kimutai Korir, Naomi E. Levin, William Mace, Anthony N. Macharia, Jay Quade, and Christopher H. Remien. 2011. \u201cWoody Cover and Hominin Environments in the Past 6 Million Years.\u201d <em>Nature<\/em> 476, no. 7358 (2011): 51-56..<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clarke, Ronald J. 1998. \u201cFirst Ever Discovery of a Well-Preserved Skull and Associated Skeleton of <em>Australopithecus<\/em>.\u201d <em>South African Journal of Science<\/em> 94 (10): 460\u2013463.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Clarke, Ronald J. 2013. \u201cAustralopithecus from Sterkfontein Caves, South Africa.\u201d In <em>The Paleobiology of Australopithecus<\/em>, edited by K. E. Reed, J. G. Fleagle, and R. E. Leakey, 105\u2013123. Netherlands: Springer.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clarke, Ronald J., and Kathleen Kuman. 2019. \u201cThe Skull of StW 573, a 3.67 Ma Australopithecus Prometheus Skeleton from Sterkfontein Caves, South Africa.\u201d <em>Journal of <\/em><em>H<\/em><em>uman <\/em><em>E<\/em><em>volution<\/em> 134: 102634.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clarke, R. J., and P. V. Tobias. 1995. \u201cSterkfontein Member 2 Foot Bones of the Oldest South African Hominid.\u201d <em>Science<\/em> 269 (5223): 521\u2013524.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Constantino, P. J., and B. A. Wood. 2004. \u201cParanthropus Paleobiology\u201d. In <em>Miscelanea en <\/em><em>Homenae<\/em><em> a Emiliano Aguirre<\/em><em>,<\/em> <em>v<\/em><em>olumen III: Paleoantropologia<\/em>, edited by E. G. P\u00e9rez and S. R. Jara, 136\u2013151. Alcal\u00e1 de Henares: Museo Arqueologico Regional.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Constantino, P. J., and B. A. Wood. 2007. \u201cThe Evolution of Zinjanthropus boisei.\u201d <em>Evolutionary Anthropology: <\/em><em>Issues, News, and Reviews<\/em> 16 (2): 49\u201362.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Dart, Raymond A. 1925. \u201cAustralopithecus africanus, the Man-Ape of South Africa.\u201d <em>Nature<\/em> 115: 195\u2013199.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Darwin, Charles. 1871. <em>The Descent of Man: And Selection in Relation to Sex<\/em>. London: J. Murray.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Daver, Guillaume, F. Guy, Hassane Ta\u00efsso Mackaye, Andossa Likius, J-R. Boisserie, Abderamane Moussa, Laurent Pallas, Patrick Vignaud, and N\u00e9koulnang D. Clarisse. 2022. \"Postcranial Evidence of Late Miocene Hominin Bipedalism in Chad.\" <em>Nature<\/em> 609 (7925): 94\u2013100.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Heinzelin, Jean de, J. Desmond Clark, Tim White, William Hart, Paul Renne, Giday WoldeGabriel, Yonas Beyene, and Elisabeth Vrba. 1999. \u201cEnvironment and Behavior of 2.5-Million-Year-Old Bouri Hominids.\u201d <em>Science<\/em> 284 (5414): 625\u2013629.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">DeMenocal, Peter B. D. 2004. \u201cAfrican Climate Change and Faunal Evolution during the Pliocene\u2013Pleistocene.\u201d <em>Earth and Planetary Science Letters<\/em> 220 (1\u20132): 3\u201324.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">DeMenocal, Peter B. D. and J. Bloemendal, J. 1995. \u201cPlio-Pleistocene Climatic Variability in Subtropical Africa and the Paleoenvironment of Hominid Evolution: A Combined Data-Model Approach.\u201d In <em>Paleoclimate and Evolution, with Emphasis on Human Origins<\/em>, edited by E. S. Vrba, G. H. Denton, T. C. Partridge, and L. H. Burckle, 262\u2013288. New Haven: Yale University Press.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Dirks, Paul HGM, Job M. Kibii, Brian F. Kuhn, Christine Steininger, Steven E. Churchill, Jan D. Kramers, Robyn Pickering, Daniel L. Farber, Anne-Sophie M\u00e9riaux, Andy I. R. Herries, Geoffrey C. P. King, And Lee R. Berger. 2010. \u201cGeological Setting and Age of <em>Australopithecus sediba<\/em> from Southern Africa.\u201d <em>Science<\/em> 328 (5975): 205\u2013208.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Faith, J. Tyler, and Anna K. Behrensmeyer. 2013. \u201cClimate Change and Faunal Turnover: Testing the Mechanics of the Turnover-Pulse Hypothesis with South African Fossil Data.\u201d <em>Paleobiology<\/em> 39 (4): 609\u2013627.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Grine, Frederick E. 1988. \u201cNew Craniodental Fossils of <em>Paranthropus<\/em> from the Swartkrans Formation and Their Significance in \u2018Robust\u2019 Australopithecine Evolution.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by F. E. Grine, 223\u2013243. New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Grine, Frederick E., Carrie S. Mongle, John G. Fleagle, and Ashley S. Hammond. 2022. \"The Taxonomic Attribution of African Hominin Postcrania from the Miocene through the Pleistocene: Associations and Assumptions.\" <em>Journal of Human Evolution<\/em> 173: 103255.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Haile-Selassie, Yohannes, Luis Gibert, Stephanie M. Melillo, Timothy M. Ryan, Mulugeta Alene, Alan Deino, Naomi E. Levin, Gary Scott, and Beverly Z. Saylor. 2015. \u201cNew Species from Ethiopia Further Expands Middle Pliocene Hominin Diversity.\u201d <em>Nature<\/em> 521 (7553): 432\u2013433.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Haile-Selassie, Yohannes, Stephanie M. Melillo, Antonino Vazzana, Stefano Benazzi, and Timothy M. Ryan. 2019. \u201cA 3.8-Million-Year-Old Hominin Cranium from Woranso-Mille, Ethiopia.\u201d <em>Nature<\/em> 573 (7773): 214-219.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Harmand, Sonia, Jason E. Lewis, Craig S. Feibel, Christopher J. Lepre, Sandrine Prat, Arnaud Lenoble, Xavier Bo\u00ebs et al. 2015. \u201c3.3-Million-Year-Old Stone Tools from Lomekwi3, West Turkana, Kenya.\u201d <em>Nature<\/em> 521 (7552): 310\u2013316.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Hay, Richard L. 1990. \u201cOlduvai Gorge: A Case History in the Interpretation of Hominid Paleoenvironments.\u201d In <em>East Africa: Establishment of a Geologic Framework for Paleoanthropology<\/em>, edited by L. Laporte, 23\u201337<em>.<\/em> Boulder: Geological Society of America.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Hay, Richard L., and Mary D. Leakey. 1982. \u201cThe Fossil Footprints of Laetoli.\u201d <em>Scientific American<\/em> 246 (2): 50\u201357.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Hlazo, Nomawethu. 2015. \u201cParanthropus: Variation in Cranial Morphology.\u201d Honours thesis, Archaeology Department, University of Cape Town, Cape Town.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Hlazo, Nomawethu. 2018. \u201cVariation and the Evolutionary Drivers of Diversity in the Genus <em>Paranthropus<\/em>.\u201d Master\u2019s thesis, Archaeology Department, University of Cape Town, Cape Town.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Johanson, D. C., T. D. White, and Y. Coppens. 1978. \u201cA New Species of the Genus <em>Australopithecus<\/em> (Primates: Hominidae) from the Pliocene of East Africa.\u201d <em>Kirtlandia<\/em> 28: 1\u201314.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kimbel, William H. 2015. \u201cThe Species and Diversity of Australopiths.\u201d In <em>Handbook of Paleoanthropology<\/em>, 2nd ed., edited by T. Hardt, 2071\u20132105. Berlin: Springer.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kimbel, William H., and Lucas K. Delezene. 2009. \u201c\u2018Lucy\u2019 Redux: A Review of Research on <em>Australopithecus afarensis<\/em>.\u201d <em>American <\/em><em>J<\/em><em>ournal of <\/em><em>P<\/em><em>hysical <\/em><em>A<\/em><em>nthropology<\/em> 140 (S49): 2\u201348.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Kingston, John D. 2007. \u201cShifting Adaptive Landscapes: Progress and Challenges in Reconstructing Early Hominid Environments.\u201d <em>American Journal of Physical Anthropology<\/em> 134 (S45): 20\u201358.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Kingston, John D., and Terry Harrison. 2007. \u201cIsotopic Dietary Reconstructions of Pliocene Herbivores at Laetoli: Implications for Early Hominin Paleoecology.\u201d <em>Palaeogeography, Palaeoclimatology, Palaeoecology<\/em> 243 (3\u20134): 272\u2013306.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Leakey, Louis S. B. 1959. \u201cA New Fossil Skull from Olduvai.\u201d <em>Nature<\/em> 184 (4685): 491\u2013493.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Leakey, Mary 1971. <em>Olduvai Gorge<\/em>, Vol. 3. Cambridge: Cambridge University Press.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Leakey, Mary D., and Richard L. Hay. 1979. \u201cPliocene Footprints in the Laetoli Beds at Laetoli, Northern Tanzania.\u201d <em>Nature<\/em> 278 (5702): 317\u2013323.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Leakey, Meave G., Craig S. Feibel, Ian McDougall, and Alan Walker. 1995. \u201cNew Four\u2013Million-Year-Old Hominid Species from Kanapoi and Allia Bay, Kenya.\u201d <em>Nature<\/em> 376 (6541): 565\u2013571.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Meave G., Fred Spoor, Frank H. Brown, Patrick N. Gathogo, Christopher Kiarie, Louise N. Leakey, and Ian McDougall. 2001. \u201cNew Hominin Genus from Eastern Africa Shows Diverse Middle Pliocene Lineages.\u201d <em>Nature<\/em> 410 (6827): 433\u2013440.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lebatard, Anne-Elisabeth, Didier L. Bourl\u00e8s, Philippe Duringer, Marc Jolivet, R\u00e9gis Braucher, Julien Carcaillet, Mathieu Schuster et al. 2008. \u201cCosmogenic Nuclide Dating of <em>Sahelanthropus tchadensis<\/em> and <em>Australopithecus bahrelghazali<\/em>: Mio-Pliocene Hominids from Chad.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 105 (9): 3226\u20133231.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lee-Thorp, Julia. 2011. \u201cThe Demise of \u2018Nutcracker Man.\u2019\u201d <em>Proceedings of the National Academy of Sciences<\/em> 108 (23): 9319\u20139320.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lombard, Marlize, L. Y. N. Wadley, Janette Deacon, Sarah Wurz, Isabelle Parsons, Moleboheng Mohapi, Joane Swart, and Peter Mitchell. 2012. \u201cSouth African and Lesotho Stone Age Sequence Updated.\u201d <em>The South African Archaeological Bulletin<\/em> 67 (195): 123\u2013144.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Maslin, Mark A., Chris M. Brierley, Alice M. Milner, Susanne Shultz, Martin H. Trauth, and Katy E. Wilson. 2014. \u201cEast African Climate Pulses and Early Human Evolution.\u201d <em>Quaternary Science Reviews<\/em> 101: 1\u201317.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">McHenry, Henry M. 2009. \u201cHuman Evolution.\u201d In <em>Evolution: The First Four Billion Years<\/em>, edited by M. Ruse and J. Travis, 256\u2013280. Cambridge: The Belknap Press of Harvard University Press..<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Patterson, Bryan, and William W. Howells. 1967. \u201cHominid Humeral Fragment from Early Pleistocene of Northwestern Kenya.\u201d <em>Science<\/em> 156 (3771): 64\u201366.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Pickering, Robyn, and Jan D. Kramers. 2010. \u201cRe-appraisal of the Stratigraphy and Determination of New U-Pb Dates for the Sterkfontein Hominin Site.\u201d <em>Journal of Human Evoluti<\/em><em>on<\/em> 59 (1): 70\u201386.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Potts, Richard. 1998. \u201cEnvironmental Hypotheses of Hominin Evolution.\u201d <em>American Journal of Physical Anthropology<\/em> 107 (S27): 93\u2013136.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Potts, Richard. 2013. \u201cHominin Evolution in Settings of Strong Environmental Variability.\u201d <em>Quaternary Science Reviews<\/em> 73: 1\u201313.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Rak, Yoel. 1983. <em>The <\/em><em>A<\/em><em>ustralopithecine <\/em><em>F<\/em><em>ace<\/em>. New York: Academic Press.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Rak, Yoel. 1988. \u201cOn Variation in the Masticatory System of <em>Australopithecus boisei<\/em>.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by M. Ruse and J. Travis, 193\u2013198<em>.<\/em> New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Semaw, Sileshi. 2000. \u201cThe World\u2019s Oldest Stone Artefacts from Gona, Ethiopia: Their Implications for Understanding Stone Technology and Patterns of Human Evolution between 2.6 Million Years Ago and 1.5 Million Years Ago.\u201d <em>Journal of Archaeological Science<\/em> 27(12): 1197\u20131214.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Shipman, Pat. 2002. <em>The Man Who Found the Missing Link: Eug<\/em><em>e<\/em><em>ne Dubois and <\/em><em>h<\/em><em>is Lifelong Quest to Prove Darwin Right<\/em>. New York: Simon &amp; Schuster.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Spoor, Fred. 2015. \u201cPalaeoanthropology: The Middle Pliocene Gets Crowded.\u201d<em> Nature<\/em> 521 (7553): 432\u2013433.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Strait, David S., Frederick E. Grine, and Marc A. Moniz. 1997. A Reappraisal of Early Hominid Phylogeny.\u201d <em>Journal of <\/em><em>H<\/em><em>uman <\/em><em>E<\/em><em>volution<\/em> 32 (1): 17\u201382.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Thackeray, J. Francis. 2000. \u201c\u2018Mrs. Ples\u2019 from Sterkfontein: Small Male or Large Female?\u201d <em>The South African Archaeological <\/em><em>Bulletin<\/em> 55: 155\u2013158.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Thackeray, J. Francis, Jos\u00e9 Braga, Jacques Treil, N. Niksch, and J. H. Labuschagne. 2002. \u201c\u2018Mrs. Ples\u2019 (Sts 5) from Sterkfontein: An Adolescent Male?\u201d <em>South African Journal of Science<\/em> 98 (1\u20132): 21\u201322.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Toth, Nicholas. 1985. \u201cThe Oldowan Reassessed.\u201d <em>Journal of Archaeological Science<\/em>\u00a012 (2): 101\u2013120.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Vrba, E. S. 1988. \u201cLate Pliocene Climatic Events and Hominid Evolution.\u201d In <em>The <\/em><em>E<\/em><em>volutionary <\/em><em>H<\/em><em>istory of the <\/em><em>R<\/em><em>obust Australopithecines<\/em>, edited by F. E. Grine, 405\u2013426. New York: Aldine.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Vrba, Elisabeth S. 1998. \u201cMultiphasic Growth Models and the Evolution of Prolonged Growth Exemplified by Human Brain Evolution.\u201d <em>Journal of Theoretical Biology<\/em> 190 (3): 227\u2013239.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Vrba, Elisabeth S. 2000. \u201cMajor Features of Neogene Mammalian Evolution in Africa.\u201d In <em>Cenozoic <\/em><em>G<\/em><em>eology of <\/em><em>S<\/em><em>outhern Africa<\/em>, edited by T. C. Partridge and R. Maud, 277\u2013304<em>.<\/em> Oxford: Oxford University Press.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Walker, Alan C., and Richard E. Leakey. 1988. \u201cThe Evolution of <em>Australopithecus boisei<\/em>.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by F. E. Grine, 247\u2013258. New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Walker, Alan, Richard E. Leakey, John M. Harris, and Francis H. Brown. 1986. \u201c2.5-my <em>Australopithecus boisei<\/em> from West of Lake Turkana, Kenya.\u201d <em>Nature<\/em> 322 (6079): 517\u2013522.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Ward, Carol, Meave Leakey, and Alan Walker. 1999. \u201cThe New Hominid Species <em>Australopithecus anamensis<\/em>.\u201d <em>Evolutionary Anthropology<\/em> 7 (6): 197\u2013205.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">White, Tim D. 1988. \u201cThe Comparative Biology of \u2018Robust\u2019 Australopithecus: Clues from Content.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by F. E. Grine, 449\u2013483. New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">White, Tim D., Gen Suwa, and Berhane Asfaw. 1994. \u201c<em>Australopithecus ramidus<\/em>, a New Species of Early Hominid from Aramis, Ethiopia.\u201d <em>Nature<\/em> 371 (6495): 306\u2013312.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Wood, Bernard. 2010. \u201cReconstructing Human Evolution: Achievements, Challenges, and Opportunities.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 10 (2): 8902\u20138909.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Wood, Bernard, and Eve K. Boyle. 2016. \u201cHominin Taxic Diversity: Fact or Fantasy?\u201d <em>Yearbook of Physical Anthropology<\/em> 159 (S61): 37\u201378.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Wood, Bernard, and Kes Schroer. 2017. \u201cParanthropus: Where Do Things Stand?\u201d In <em>Human Paleontology and Prehistory<\/em>, edited by A. Marom and E. Hovers, 95\u2013107. New York: Springer, Cham.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Acknowledgements<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">All of the authors in this section are students and early career researchers in paleoanthropology and related fields in South Africa (or at least have worked in South Africa). We wish to thank everyone who supports young and diverse talent in this field and would love to further acknowledge Black, African, and female academics who have helped pave the way for us.<\/span><\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1764\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1764\"><div tabindex=\"-1\"><p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kerryn Warren, Ph.D., Grad Coach International<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lindsay Hunter, M.A., University of Iowa<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Navashni Naidoo, M.Sc., University of Cape Town<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Silindokuhle Mavuso, M.Sc., University of Witwatersrand<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\"><em>This chapter is a revision from <\/em><em>\"<\/em><a class=\"rId7\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\"><em>Chapter 9: Early Hominins<\/em><\/a><em>\" <\/em><em>by Kerryn Warren, K. Lindsay Hunter, Navashni Naidoo, Silindokuhle Mavuso, Kimberleigh Tommy, Rosa Moll, and Nomawethu Hlazo<\/em><em>. In <\/em><a class=\"rId8\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, first edition<\/em><\/a><em>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under <\/em><a class=\"rId9\" style=\"color: #000000\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\"><em>CC BY-NC 4.0<\/em><\/a><em>. <\/em><\/span><\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\"><span style=\"color: #000000\"><span style=\"color: #ffffff\">Learning Objectives<\/span><br \/>\n<\/span><\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li><span style=\"color: #000000\">Understand what is meant by \u201cderived\u201d and \u201cancestral\u201d traits and why this is relevant for understanding early hominin evolution.<\/span><\/li>\n<li><span style=\"color: #000000\">Understand changing paleoclimates and paleoenvironments as potential factors influencing early hominin adaptations.<\/span><\/li>\n<li><span style=\"color: #000000\">Describe the anatomical changes associated with bipedalism and dentition in early hominins, as well as their implications..<\/span><\/li>\n<li><span style=\"color: #000000\">Describe early hominin genera and species, including their currently understood dates and geographic expanses.<\/span><\/li>\n<li><span style=\"color: #000000\">Describe the earliest stone tool techno-complexes and their impact on the transition from early hominins to our genus.<\/span><\/li>\n<\/ul>\n<\/div>\n<\/div>\n<h2 class=\"__UNKNOWN__\"><span style=\"color: #000000\">Defining Hominins<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">It is through our study of our hominin ancestors and relatives that we are exposed to a world of \u201cmight have beens\u201d: of other paths not taken by our species, other ways of being human. But to better understand these different evolutionary trajectories, we must first define the terms we are using. If an imaginary line were drawn between ourselves and our closest relatives, the great apes, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1200\">bipedalism<\/a><\/strong> (or habitually walking upright on two feet) is where that line would be. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1270\">Hominin<\/a><\/strong>, then, means everyone on \u201cour\u201d side of the line: humans and all of our extinct bipedal ancestors and relatives since our divergence from the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1272\">last common ancestor (LCA)<\/a><\/strong> we share with chimpanzees.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Historic interpretations of our evolution, prior to our finding of early hominin <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1274\"><strong>fossils<\/strong><\/a>, varied. Debates in the mid-1800s regarding hominin origins focused on two key issues:<\/span><\/p>\n<ul>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">Where did we evolve?<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">Which traits evolved first?<\/span><\/li>\n<\/ul>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff00ff\">Charles Darwin hypothesized that we evolved in Africa, as he was convinced that we shared greater commonality with chimpanzees and gorillas on the continent (Darwin 1871). Others, such as Ernst Haeckel and Eug\u00e8ne Dubois, insisted that we were closer in affinity to orangutans and that we evolved in Eurasia where, until the discovery of the Taung Child in South Africa in 1924, all humanlike fossils (of Neanderthals and <em>Homo erectus<\/em>) had been found (Shipman 2002).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Within this conversation, naturalists and early <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1276\">paleoanthropologists<\/a><\/strong> (people who study human evolution) speculated about which human traits came first. These included the evolution of a big brain (<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1278\"><strong>encephalization<\/strong><\/a>), the evolution of the way in which we move about on two legs (bipedalism), and the evolution of our flat faces and small teeth (indications of dietary change). Original hypotheses suggested that, in order to be motivated to change diet and move about in a bipedal fashion, the large brain needed to have evolved first, as is seen in the fossil species mentioned above.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">However, we now know that bipedal locomotion is one of the first things that evolved in our lineage, with early relatives having more apelike dentition and small brain sizes. While brain size expansion is seen primarily in our genus, <em>Homo<\/em>, earlier hominin brain sizes were highly variable between and within taxa, from 300 cc (cranial capacity, cm<sup>3<\/sup>), estimated in <em>Ardipithecus<\/em>, to 550 cc, estimated in <em>Paranthropus boisei<\/em>. The lower estimates are well within the range of variation of nonhuman extant great apes. In addition, body size variability also plays a role in the interpretation of whether brain size could be considered large or small for a particular species or specimen. In this chapter, we will tease out the details of early hominin evolution in terms of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1280\"><strong>morphology<\/strong><\/a> (i.e. the study of the form, size, or shape of things; in this case, skeletal parts).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">We also know that early human evolution occurred in a very complicated fashion. There were multiple species (multiple genera) that featured diversity in their diets and locomotion. Specimens have been found all along the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1282\"><strong>East African Rift System <\/strong>(<strong>EARS)<\/strong><\/a>; that is, in Ethiopia, Kenya, Tanzania, and Malawi; see Figure 9.1), in limestone caves in South Africa, and in Chad. Dates of these early relatives range from around 7 million years ago (mya) to around 1 mya, overlapping temporally with members of our genus, <em>Homo<\/em>.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 610px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/06\/image38.png\" alt=\"Patchy green mountain alongside a deep sandy valley in East Africa.\" width=\"610\" height=\"277\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.1: East African Rift System (EARS). Credit: <a href=\"https:\/\/www.flickr.com\/photos\/ninara\/8624605781\/in\/photolist-x2yH7-x2yHe-VfVWuD-e98mPF-SzzjsU-2bsBZhC-2hHec7m-xtJ7Ez-NXnXvh-7Yg3uo-2cS3FgG-2hjo1Dc-2hjGoTS-nnumi8-82U66W-dMNn7B-8jdVbd-NWDg8-NW6fj-ebhx5w-bkFv1G-Ct5ZD-5JQk8A-y6TgAc-x9k6oe-2ebLTDC-WcPMnJ-2ekh6CS-Cu3LH-xNHDFK-9RUsZi-94jVt4-P46uiB-QFyjyE-crU8N7-5JLJKV-2ekSgk8-5JL454-2cPgZrF-2bHfQZu-dMTVPN-6yUbeN-jzMicQ-48XjU9-2etR2Ze-Styrvw-crU7V7-2wakq3-crU6Z1-2etR2XR\/\">IMG_1696 Great Rift Valley<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/ninara\/\">Ninara<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\">CC BY 2.0 License<\/a>.<\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Yet there is still so much to understand. Modern debates now look at the relatedness of these species to us and to one another, and they consider which of these species were able to make and use tools. As a result, every <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1284\">site<\/a><\/strong> discovery in the patchy hominin fossil record tells us more about our evolution. In addition, recent scientific techniques (not available even ten years ago) provide new insights into the diets, environments, and lifestyles of these ancient relatives.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In the past, <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1288\"><strong>taxonom<\/strong><strong>y<\/strong><\/a> was primarily based on morphology. Today it is tied to known relationships based on molecular <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1290\"><strong>phylogeny<\/strong><\/a> (e.g., based on DNA) or a combination of the two. This is complicated when applied to living <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1286\"><strong>taxa<\/strong><\/a>, but becomes much more difficult when we try to categorize ancestor-descendant relationships for long-extinct species whose molecular information is no longer preserved. We therefore find ourselves falling back on morphological comparisons, often of teeth and partially fossilized skeletal material.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">It is here that we turn to the related concepts of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1292\">cladistics<\/a> <\/strong>and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1294\">phylogenetics<\/a><\/strong><strong>. <\/strong><span style=\"background-color: #ff99cc\">Cladistics groups organisms according to their last common ancestors based on shared <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1126\">derived traits<\/a><\/strong>. <\/span>In the case of early hominins, these are often morphological traits that differ from those seen in earlier populations. These new or modified traits provide evidence of evolutionary relationships, and organisms with the same derived traits are grouped in the same <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1116\">clade<\/a> <\/strong>(Figure 9.2). <span style=\"background-color: #ff99cc\">For example, if we use feathers as a trait, we can group pigeons and ostriches into the clade of birds.<\/span> In this chapter, we will examine the grouping of the Robust Australopithecines, whose cranial and dental features differ from those of earlier hominins, and therefore are considered derived.<\/span><\/p>\n<figure style=\"width: 708px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image48.png\" alt=\"Phylogenetic tree shows clades and non clade groupings.\" width=\"708\" height=\"192\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.2: Clades refer to groups of species or taxa that share a common ancestor. In <span class=\"ILfuVd\" lang=\"en\"><span class=\"hgKElc\">a phylogeny, a clade is a complete group of lineages, including their last common ancestor. Groupings that do not include a common ancestor and <em>all<\/em> of its descendants are not clades. <\/span><\/span>Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\">Clades (Figure 9.2)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Katie Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Dig Deeper: Problems Defining Hominin Species<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000\">It is worth noting that species designations for early hominin specimens are often highly contested. This is due to the fragmentary nature of the fossil record, the large timescale (millions of years) with which paleoanthropologists need to work, and the difficulty in evaluating whether morphological differences and similarities are due to meaningful phylogenetic or biological differences or subtle differences\/variation in niche occupation or time. In other words, do morphological differences really indicate different species? How would classifying species in the paleoanthropological record compare with classifying living species today, for whom we can sequence genomes and observe lifestyles?<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">There are also broader philosophical differences among researchers when it comes to paleo-species designations. Some scientists, known as \u201c<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1296\"><strong>lumpers<\/strong>,<\/a>\u201d argue that large variability is expected among multiple populations in a given species over time. These researchers will therefore prefer to \u201clump\u201d specimens of subtle differences into single taxa. Others, known as \u201c<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1298\"><strong>splitters<\/strong><\/a>,\u201d argue that species variability can be measured and that even subtle differences can imply differences in niche occupation that are extreme enough to mirror modern species differences. In general, splitters would consider geographic differences among populations as meaning that a species is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1300\">polytypic<\/a><\/strong> <span style=\"background-color: #ff99cc\">(i.e., capable of interacting and breeding biologically but having morphological population differences)<\/span>. This is worth keeping in mind when learning about why species designations may be contested.<\/span><\/p>\n<figure style=\"width: 520px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image8.jpg\" alt=\"A graph shows a curved line depicting changes in morphology among two species over time.\" width=\"520\" height=\"292\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.3: This graph demonstrates the concept of a chronospecies, where one species (Species A) \u201cevolves\u201d into another (Species B). Credit: Chronospecies original to Explorations: An Open Invitation to Biological Anthropology, 2nd edition by Kerryn Warren is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\"><span style=\"color: #000000\">This further plays a role in evaluating ancestry. Debates over which species \u201cgave rise\u201d to which continue to this day. It is common to try to create \u201clineages\u201d of species to determine when one species evolved into another over time. We refer to these as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1302\"><strong>chronospecies<\/strong><\/a> (Figure 9.3). Constructed hominin phylogenetic trees are routinely variable, changing with new specimen discoveries, new techniques for evaluating and comparing species, and, some have argued, nationalist or biased interpretations of the record. More recently, some researchers have shifted away from \u201ctreelike\u201d models of ancestry toward more nuanced metaphors such as the \u201cbraided stream,\u201d where some levels of interbreeding among species and populations are seen as natural processes of evolution.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Finally, it is worth considering the process of fossil discovery and publication. Some fossils are easily diagnostic to a species level and allow for easy and accurate interpretation. Some, however, are more controversial. This could be because they do not easily preserve or are incomplete, making it difficult to compare and place within a specific species (e.g., a fossil of a patella or knee bone). Researchers often need to make several important claims when announcing or publishing a find: a secure date (if possible), clear association with other finds, and an adequate comparison among multiple species (both extant and fossil). Therefore, it is not uncommon that an important find was made years before it is scientifically published.<\/span><\/p>\n<\/div>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Paleoenvironment and Hominin Evolution<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There is no doubt that one of the major selective pressures in hominin evolution is the environment. <span style=\"background-color: #ff9900\"><span style=\"background-color: #ffff00\">Large-scale changes in global and regional climate, as well as alterations to the environment, are<\/span><span style=\"background-color: #ffff00\"> (<\/span><\/span><em style=\"background-color: transparent\"><span style=\"text-decoration: underline\">thought to be<\/span>)<\/em><span style=\"background-color: #ffff00\"><del> all<\/del> linked to (all) hominin diversification, dispersal, and extinction<\/span> (Maslin et al. 2014). Environmental reconstructions often use modern analogues. Let us take, for instance, the hippopotamus. It is an animal that thrives in environments that have abundant water to keep its skin cool and moist. If the environment for some reason becomes drier, it is expected that hippopotamus populations will reduce. If a drier environment becomes wetter, it is possible that hippopotamus populations may be attracted to the new environment and thrive. Such instances have occurred multiple times in the past, and the bones of some <strong style=\"background-color: transparent\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1308\">fauna<\/a><\/strong> (i.e., animals, like the hippopotamus) that are sensitive to these changes give us insights into these events.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Yet reconstructing a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1304\">paleoenvironment<\/a><\/strong> relies on a range of techniques, which vary depending on whether research interests focus on local changes or more global environmental changes\/reconstructions. For local environments (such as a single site or region), comparing the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1306\">faunal assemblages<\/a> <\/strong>(collections of fossils of animals found at a site) with animals found in certain modern environments allows us to determine if past environments mirror current ones in the region. Changes in the faunal assemblages, as well as when they occur and how they occur, tell us about past environmental changes. Other techniques are also useful in this regard. Chemical analyses, for instance, can reveal the diets of individual fauna, providing clues as to the relative wetness or dryness of their environment (e.g., nitrogen <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1310\"><strong>isotopes<\/strong><\/a>; Kingston and Harrison 2007).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Global climatic changes in the distant past, which fluctuated between being colder and drier and warmer and wetter on average, would have global implications for environmental change (Figure 9.4). These can be studied by comparing marine core and terrestrial soil data across multiple sites. These techniques are based on chemical analysis, such as examination of the nitrogen and oxygen isotopes in shells and sediments. Similarly, analyzing pollen grains shows which kinds of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1312\"><strong>flora<\/strong> <\/a> survived in an environment at a specific time period. There are multiple lines of evidence that allow us to visualize global climate trends over millions of years (although it should be noted that the direction and extent of these changes could differ by geographic region).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 649px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image12-1-1.png\" alt=\"Chart shows cyclical carbon dioxide levels from 800,000 years ago until today.\" width=\"649\" height=\"406\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.4: This graph, based on the comparison of atmospheric samples contained in ice cores and more recent direct measurements, illustrates how atmospheric CO\u2082 has fluctuated over time and increased sharply since the Industrial Revolution. The graph also shows that since 800,000ya (and before) atmospheric CO\u2082 has never exceeded 300 parts per million (ppm). In 1950 it was 310ppm. Today atmospheric CO\u2082 has spiked to over 410 ppm. Credit: <a href=\"https:\/\/climate.nasa.gov\/evidence\/\">CO\u2082 increase since the Industrial Revolution<\/a> by <a href=\"https:\/\/www.nasa.gov\/\">NASA<\/a> is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a> and is used within <a href=\"https:\/\/www.nasa.gov\/multimedia\/guidelines\/index.html\">NASA guidelines on re-use<\/a>. Original from Luthi, D., et al.. 2008; Etheridge, D.M., et al. 2010; Vostok ice core data\/J.R. Petit et al.; NOAA Mauna Loa CO<a href=\"https:\/\/climate.nasa.gov\/evidence\/\">\u2082<\/a> record..<\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Both local and global climatic\/environmental changes have been used to understand factors affecting our evolution (DeHeinzelin et al. 1999; Kingston 2007). Environmental change acts as an important factor regarding the onset of several important hominin traits seen in early hominins and discussed in this chapter. Namely, the environment has been interpreted as the following:<\/span><\/p>\n<ul>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">the driving force behind the evolution of bipedalism,<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">the reason for change and variation in early hominin diets, and<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">the diversification of multiple early hominin species.<\/span><\/li>\n<\/ul>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are numerous hypotheses regarding how climate has driven and continues to drive human evolution. Here, we will focus on just three popular hypotheses.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Savannah Hypothesis (or Aridity Hypothesis)<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The hypothesis:<\/strong> This popular theory suggests that the expansion of the savannah (or less densely forested, drier environments) forced early hominins from an <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1314\"><strong>arboreal<\/strong> <\/a> lifestyle (one living in trees) to a terrestrial one where bipedalism was a more efficient form of locomotion (Figure 9.5). It was first proposed by Darwin (1871) and supported by anthropologists like Raymond Dart (1925). However, this idea was supported by little fossil or paleoenvironmental evidence and was later refined as the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1318\">Aridity Hypothesis<\/a><\/strong>. This hypothesis states that the long-term <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1316\">aridification<\/a><\/strong> and, thereby, expansion of savannah biomes were drivers in diversification in early hominin evolution (deMenocal 2004; deMenocal and Bloemendal 1995). It advocates for periods of accelerated aridification leading to early hominin speciation events.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 647px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image46.png\" alt=\"Photograph showing a dry, open savannah environment.\" width=\"647\" height=\"486\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.5: The African savannah grew during early hominin evolution. This may have forced early hominins from an arboreal lifestyle to a terrestrial one, where bipedalism was a more efficient form of locomotion. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:African_savannah_@_Masai_Mara_(21308330314).jpg\">African savannah @ Masai Mara (21308330314)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/132394214@N04\">Leo Li<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\">CC BY 2.0 License<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The evidence:<\/strong> While early bipedal hominins are often associated with wetter, more closed environments (i.e., not the Savannah Hypothesis), both marine and terrestrial records seem to support general cooling, drying conditions, with isotopic records indicating an increase in grasslands (i.e., colder and wetter climatic conditions) between 8 mya and 6 mya across the African continent (Cerling et al. 2011). This can be contrasted with later climatic changes derived from aeolian dust records (sediments transported to the site of interest by wind), which demonstrate increases in seasonal rainfall between 3 mya and 2.6 mya, 1.8 mya and 1.6 mya, and 1.2 mya and 0.8 mya (deMenocal 2004; deMenocal and Bloemendal 1995).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Interpretation(s):<\/strong> Despite a relatively scarce early hominin record, it is clear that two important factors occur around the time period in which we see increasing aridity. The first factor is the diversification of taxa, where high morphological variation between specimens has led to the naming of multiple hominin genera and species. The second factor is the observation that the earliest hominin fossils appear to have traits associated with bipedalism and are dated to around the drying period (as based on isotopic records). Some have argued that it is more accurately a combination of bipedalism and arboreal locomotion, which will be discussed later. However, the local environments in which these early specimens are found (as based on the faunal assemblages) do not appear to have been dry.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Turnover Pulse Hypothesis<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The hypothesis:<\/strong> In 1985, paleontologist Elisabeth Vbra noticed that in periods of extreme and rapid climate change, <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1326\"><strong>ungulates<\/strong><\/a> (hoofed mammals of various kinds) that had generalized diets fared better than those with specialized diets (Vrba 1988, 1998). <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1322\"><strong>Specialist<\/strong><\/a> eaters <span style=\"background-color: #ff99cc\">(those who rely primarily on specific food types)<\/span> faced extinction at greater rates than their <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1320\">generalist<\/a> <\/strong><span style=\"background-color: #ff99cc\">(those who can eat more varied and variable diets) <\/span>counterparts because they were unable to adapt to new environments (Vrba 2000). Thus, periods with extreme climate change would be associated with high <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1324\"><strong>faunal turnover<\/strong><\/a>: that is, the extinction of many species and the speciation, diversification, and migration of many others to occupy various niches.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The evidence:<\/strong> The onset of the<strong> <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1400\">Quaternary Ice Age<\/a><\/strong>, between 2.5 mya and 3 mya, brought extreme global, cyclical <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1328\"><strong>interglacial<\/strong> <\/a> and <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1330\"><strong>glacial<\/strong><\/a> periods (warmer, wetter periods with less ice at the poles, and colder, drier periods with more ice near the poles). Faunal evidence from the Turkana basin in East Africa indicates multiple instances of faunal turnover and extinction events, in which global climatic change resulted in changes from closed\/forested to open\/grassier habitats at single sites (Behrensmeyer et al. 1997; Bobe and Behrensmeyer 2004). Similarly, work in the Cape Floristic Belt of South Africa shows that extreme changes in climate play a role in extinction and migration in ungulates. While this theory was originally developed for ungulates, its proponents have argued that it can be applied to hominins as well. However, the link between climate and speciation is only vaguely understood (Faith and Behrensmeyer 2013).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Interpretation(s):<\/strong> While the evidence of rapid faunal turnover among ungulates during this time period appears clear, there is still some debate around its usefulness as applied to the paleoanthropological record. Specialist hominin species do appear to exist for long periods of time during this time period, yet it is also true that <em>Homo<\/em>, a generalist genus with a varied and adaptable diet, ultimately survives the majority of these fluctuations, and the specialists appear to go extinct.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Variability Selection Hypothesis<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The hypothesis: <\/strong>This hypothesis was first articulated by paleoanthropologist Richard Potts (1998). It links the high amount of climatic variability over the last 7 million years to both behavioral and morphological changes. Unlike previous notions, this hypothesis states that hominin evolution does not respond to habitat-specific changes or to specific aridity or moisture trends. Instead, long-term environmental unpredictability over time and space influenced morphological and behavioral adaptations that would help hominins survive, regardless of environmental context (Potts 1998, 2013). The Variability Selection Hypothesis states that hominin groups would experience varying degrees of natural selection due to continually changing environments and potential group isolation. This would allow certain groups to develop genetic combinations that would increase their ability to survive in shifting environments. These populations would then have a genetic advantage over others that were forced into habitat-specific adaptations (Potts 2013).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The evidence:<\/strong> The evidence for this theory is similar to that for the Turnover Pulse Hypothesis: large climatic variability and higher survivability of generalists versus specialists. However, this hypothesis accommodates for larger time-scales of extinction and survival events.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Interpretation(s):<\/strong> In this way, the Variability Selection Hypothesis allows for a more flexible interpretation of the evolution of bipedalism in hominins and a more fluid interpretation of the Turnover Pulse Hypothesis, where species turnover is meant to be more rapid. In some ways, this hypothesis accommodates both environmental data and our interpretations of an evolution toward greater variability among species and the survivability of generalists.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000;background-color: #ff99cc\"><strong>Paleoenvironment Summary<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff99cc\">Some hypotheses presented in this section pay specific attention to habitat (Savannah Hypothesis) while others point to large-scale climatic forces (Variability Selection Hypothesis). Some may be interpreted to describe the evolution of traits such as bipedalism (Savannah Hypothesis), and others generally explain the diversification of early hominins (Turnover Pulse and Variability Selection Hypotheses). While there is no consensus as to how the environment drove our evolution, it is clear that the environment shaped both habitat and resource availability in ways that would have influenced our early ancestors physically and behaviorally.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Derived Adaptations: Bipedalism<br \/>\n<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The unique form of locomotion exhibited by modern humans, called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1332\">obligate bipedalism<\/a><\/strong>, is important in distinguishing our species from the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1334\"><strong>extant<\/strong><\/a> (living) great apes. The ability to walk habitually upright is thus considered one of the defining attributes of the hominin lineage. We also differ from other animals that walk bipedally (such as kangaroos) in that we do not have a tail to balance us as we move.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The origin of bipedalism in hominins has been debated in paleoanthropology, but at present there are two main <span style=\"background-color: #ff99cc\">ideas<\/span>: <span style=\"text-decoration: underline\">(<em>theories<\/em>)<\/span><\/span><\/p>\n<ol>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">early hominins initially lived in trees, but increasingly started living on the ground, so we were a product of an arboreal last common ancestor (LCA) or,<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">our LCA was a terrestrial quadrupedal knuckle-walking species, more similar to extant chimpanzees.<\/span><\/li>\n<\/ol>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Most research supports the first theory of an arboreal LCA based on skeletal morphology of early hominin genera that demonstrate adaptations for climbing but not for knuckle-walking. This would mean that both humans and chimpanzees can be considered \u201cderived\u201d in terms of locomotion since chimpanzees would have independently evolved knuckle-walking.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are many current ideas regarding selective pressures that would lead to early hominins adapting upright posture and locomotion. Many of these selective pressures, as we have seen in the previous section, coincide with a shift in environmental conditions, supported by paleoenvironmental data. In general, however, it appears that, like extant great apes, early hominins thrived in forested regions with dense tree coverage, which would indicate an arboreal lifestyle. As the environmental conditions changed and a savannah\/grassland environment became more widespread, the tree cover would become less dense, scattered, and sparse such that bipedalism would become more important.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are several proposed selective pressures for bipedalism:<\/span><\/p>\n<ol>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\"><strong>E<\/strong><strong>nergy conservation:<\/strong> Modern bipedal humans conserve more energy than extant chimpanzees, which are predominantly knuckle-walking quadrupeds when walking over land. While chimpanzees, for instance, are faster than humans terrestrially, they expend large amounts of energy being so. Adaptations to bipedalism include \u201cstacking\u201d the majority of the weight of the body over a small area around the center of gravity (i.e., the head is above the chest, which is above the pelvis, which is over the knees, which are above the feet). This reduces the amount of muscle needed to be engaged during locomotion to \u201cpull us up\u201d and allows us to travel longer distances expending far less energy.<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\"><strong>T<\/strong><strong>hermoregulation:<\/strong> Less surface area (i.e., only the head and shoulders) is exposed to direct sunlight during the hottest parts of the day (i.e., midday). This means that the body has less need to employ additional \u201ccooling\u201d mechanisms such as sweating, which additionally means less water loss.<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Bipedalism <\/strong><span style=\"text-decoration: underline\">(Freeing of Hands)<\/span><strong>: <\/strong>This method of locomotion freed up our ancestors\u2019 hands such that they could more easily gather food and carry tools or infants. This further enabled the use of hands for more specialized adaptations associated with the manufacturing and use of tools.<\/span><\/li>\n<\/ol>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">These selective pressures are not mutually exclusive. Bipedality could have evolved from a combination of these selective pressures, in ways that increased the chances of early hominin survival.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Skeletal Adaptations for Bipedalism<\/strong><\/span><\/h3>\n<figure style=\"width: 405px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image10-1.png\" alt=\"A full human skeleton and gorilla skeleton standing in upright positions next to each other.\" width=\"405\" height=\"452\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.6: Compared to gorillas (right) and other apes, humans (left) have highly specialized adaptations to facilitate bipedal locomotion. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Primatenskelett-drawing.jpg\">Skeleton of human (1) and gorilla (2), unnaturally sketched<\/a> by unknown from Brehms Tierleben, Small Edition 1927 is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Humans have highly specialized adaptations to facilitate obligate bipedalism (Figure 9.6). Many of these adaptations occur within the soft tissue of the body (e.g., muscles and tendons). However, when analyzing the paleoanthropological record for evidence of the emergence of bipedalism, all that remains is the fossilized bone. Interpretations of locomotion are therefore often based on comparative analyses between fossil remains and the skeletons of extant primates with known locomotor behaviors. These adaptations occur throughout the skeleton and are summarized in Figure 9.7.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The majority of these adaptations occur in the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1336\"><strong>postcranium<\/strong><\/a> <span style=\"background-color: #ff99cc\">(the skeleton from below the head)<\/span> and are outlined in Figure 9.7. In general, these adaptations allow for greater stability and strength in the lower limb, by allowing for more shock absorption, for a larger surface area for muscle attachment, and for the \u201cstacking\u201d of the skeleton directly over the center of gravity to reduce energy needed to be kept upright. These adaptations often mean less flexibility in areas such as the knee and foot.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">However, these adaptations come at a cost. Evolving from a nonobligate bipedal ancestor means that the adaptations we have are evolutionary compromises. For instance, the valgus knee (angle at the knee) is an essential adaptation to balance the body weight above the ankle during bipedal locomotion. However, the strain and shock absorption at an angled knee eventually takes its toll. For example, runners often experience joint pain. Similarly, the long neck of the femur absorbs stress and accommodates for a larger pelvis, but it is a weak point, resulting in hip replacements being commonplace among the elderly, especially in cases where the bone additionally weakens through osteoporosis. Finally, the S-shaped curve in our spine allows us to stand upright, relative to the more curved C-shaped spine of an LCA. Yet the weaknesses in the curves can lead to pinching of nerves and back pain. Since many of these problems primarily are only seen in old age, they can potentially be seen as an evolutionary compromise.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Despite relatively few postcranial fragments, the fossil record in early hominins indicates a complex pattern of emergence of bipedalism. Key features, such as a more anteriorly placed foramen magnum, are argued to be seen even in the earliest discovered hominins, indicating an upright posture (Dart 1925). Some early species appear to have a mix of ancestral (arboreal) and derived (bipedal) traits, which indicates a mixed locomotion and a more <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1402\">mosaic evolution<\/a><\/strong> of the trait. Some early hominins appear to, for instance, have bowl-shaped pelvises (hip bones) and angled femurs suitable for bipedalism but also have retained an opposable <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1338\">hallux<\/a><\/strong> (big toe) or curved fingers and longer arms (for arboreal locomotion). These mixed morphologies may indicate that earlier hominins were not fully obligate bipeds and thus thrived in mosaic environments.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Yet the associations between postcranial and the more diagnostic cranial fossils and bones are not always clear, muddying our understanding of the specific species to which fossils belong (Grine et al. 2022).<\/span><\/p>\n<div align=\"left\">\n<table class=\"grid\">\n<caption>\n<p class=\"import-Normal\" style=\"text-align: left\"><span style=\"color: #000000\">Figure 9.7: Skeletal comparisons between modern humans (obligate bipeds) and nonobligate bipeds (e.g., chimpanzees). Credit: <a class=\"rId34\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\">Skeletal comparisons between modern humans and <\/a><a class=\"rId35\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\">nonobligate<\/a><a class=\"rId36\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\"> bipeds (Figure 9.6)<\/a> original to <a class=\"rId37\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a class=\"rId38\" style=\"color: #000000\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/span><\/p>\n<\/caption>\n<thead>\n<tr>\n<td style=\"width: 97.998px\"><strong>Region<\/strong><\/td>\n<td style=\"width: 106.992px\"><strong>Feature<\/strong><\/td>\n<td style=\"width: 366.992px\"><strong>Obligate Biped (H. sapiens)<\/strong><\/td>\n<td style=\"width: 310px\"><strong>Nonobligate Biped<\/strong><\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr>\n<td style=\"width: 97.998px\">Cranium<\/td>\n<td style=\"width: 106.992px\">Position of the foramen magnum<\/td>\n<td style=\"width: 366.992px\">Positioned inferiorly (immediately under the cranium) so that the head rests on top of the vertebral column for balance and support (head is perpendicular to the ground).<\/td>\n<td style=\"width: 310px\">Posteriorly positioned (to the back of the cranium). Head is positioned parallel to the ground.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Body proportions<\/td>\n<td style=\"width: 366.992px\">Shorter upper limb (not used for locomotion).<\/td>\n<td style=\"width: 310px\">Longer upper limbs (used for locomotion).<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Spinal curvature<\/td>\n<td style=\"width: 366.992px\">S-curve due to pressure exerted on the spine from bipedalism (lumbar lordosis).<\/td>\n<td style=\"width: 310px\">C-curve.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Vertebrae<\/td>\n<td style=\"width: 366.992px\">Robust lumbar (lower-back) vertebrae (for shock absorbance and weight bearing). Lower back is more flexible than that of apes as the hips and trunk swivel when walking (weight transmission).<\/td>\n<td style=\"width: 310px\">Gracile lumbar vertebrae compared to those of modern humans.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Pelvis<\/td>\n<td style=\"width: 366.992px\">Shorter, broader, bowl-shaped pelvis (for support); very robust. Broad sacrum with large sacroiliac joint surfaces.<\/td>\n<td style=\"width: 310px\">Longer, flatter, elongated ilia; more narrow and gracile; narrower sacrum; relatively smaller sacroiliac joint surface.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Lower limb<\/td>\n<td style=\"width: 366.992px\">In general, longer, more robust lower limbs and more stable, larger joints.<\/p>\n<ul>\n<li style=\"font-weight: 400\">Large femoral head and longer neck (absorbs more stress and increases the mechanical advantage).<\/li>\n<li style=\"font-weight: 400\">Valgus knee, in which the angle of the knee positions it over the ankle and keeps the center of gravity balanced over the stance leg during stride cycle (shock absorbance).<\/li>\n<li style=\"font-weight: 400\">Distal tibia (lower leg) of humans has a large medial malleolus for stability.<\/li>\n<\/ul>\n<\/td>\n<td style=\"width: 310px\">In general, smaller, more gracile limbs with more flexible joints.<\/p>\n<ul>\n<li style=\"font-weight: 400\">Femoral neck is smaller in comparison to modern humans and shorter.<\/li>\n<li style=\"font-weight: 400\">The legs bow outward, and there is no valgus angle of the knee (no \u201cknock knees\u201d).<\/li>\n<li style=\"font-weight: 400\">The distal tibia in chimpanzees is trapezoid (wider anteriorly) for climbing and allows more flexibility.<\/li>\n<\/ul>\n<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Foot<\/td>\n<td style=\"width: 366.992px\">Rigid, robust foot, without a midtarsal break.<\/p>\n<p>Nonopposable and large, robust big toe (for push off while walking) and large heel for shock absorbance.<\/td>\n<td style=\"width: 310px\">Flexible foot, midtarsal break present (which allows primates to lift their heels independently from their feet), opposable big toe for grasping.<\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<p class=\"import-Normal\"><span style=\"color: #000000;background-color: #ccffcc\">It is also worth noting that, while not directly related to bipedalism per se, other postcranial adaptations are evident in the hominin fossil record from some of the earlier hominins. For instance, the hand and finger morphologies of many of the earliest hominins indicate adaptations consistent with arboreality. These include longer hands, more curved metacarpals and phalanges (long bones in the hand and fingers, respectively), and a shorter, relatively weaker thumb. This allows for gripping onto curved surfaces during locomotion. The earliest hominins appear to have mixed morphologies for both bipedalism and arborealism. However, among Australopiths (members of the genus, Australopithecus), there are indications for greater reliance on bipedalism as the primary form of locomotion. Similarly, adaptations consistent with tool manufacture (shorter fingers and a longer, more robust thumb, in contrast to the features associated with arborealism) have been argued to appear before the genus <em>Homo<\/em>.<\/span><\/p>\n<div class=\"textbox shaded\">\n<h2><span style=\"background-color: #00ffff\">(Special Topic with student projects: fear of snakes, cultural or biological? Biology, culture, and the fear of snakes, Snake Detection Theory)<\/span><\/h2>\n<figure style=\"width: 245px\" class=\"wp-caption alignright\"><img src=\"https:\/\/www.researchgate.net\/profile\/David-Penning\/publication\/315305069\/figure\/fig2\/AS:613947958902785@1523388008174\/Constriction-coil-postures-of-kingsnakes-and-ratsnakes-Typical-constriction-coil.png\" alt=\"Constriction coil postures of kingsnakes and ratsnakes. Typical constriction coil postures in a kingsnake, Lampropeltis getula (92 g; A), and a ratsnake, Pantherophis guttatus (86 g; B). Both snakes were constricting similarly sized mice, Mus musculus (12 g). The relative prey mass was 13% for the kingsnake and 13.9% for the ratsnake.\u00a0\" width=\"245\" height=\"245\" \/><figcaption class=\"wp-caption-text\">Constriction coil postures of kingsnakes and ratsnakes. source: https:\/\/www.researchgate.net\/figure\/Constriction-coil-postures-of-kingsnakes-and-ratsnakes-Typical-constriction-coil_fig2_315305069<\/figcaption><\/figure>\n<p>It is suggested that primates have three major predators: raptors, felines, and snakes; however, many studies show that of these carnivores, snakes were one of the first that mammals had to contend with alongside dinosaurs, as felines and raptors evolved at a much slower pace than their reptilian competition. Herpetologists trace the evolution of constricting snakes to about 100 million years ago, and by the time mammals arrived around 75 million years ago, constrictors were\u00a0 already well established as a formidable threat (Greene, 2017). \u00a0Both co-existed for millennia and each sustained selective pressures requiring them to evolve specific traits to survive. When venomous snakes eventually emerged 55 to 65 million years ago, they posed yet an additional threat to proto-primates as they required less distance for the predator to kill (2017). Alongside camouflage and silent movement techniques, it was the development of the snake\u2019s hollow fangs through which to deliver venom that was most transformative to primate evolution. As such, primates evolved their pre-conscious attention, and visual acuity to cope with this new threat; therefore, while snakes were adapting morphologically to feed themselves, they were unwittingly teaching proto-primates valuable lessons in predator detection and reacting appropriately in order to survive.<\/p>\n<p>In a 2009 Harvard University study, Lynne A. Isbell hypothesizes that envenoming snakes are linked to being directly responsible for the origins of the evolving complex brains and superior visual capacity in the lineage of anthropoids leading to humans (Isbell, 2009). Forward-facing eyes for binocular vision, depth perception, enhanced visual acuity, stereoscopic and trichromatic colour vision, all traits necessary for snake detection; and the quick motor responses from the primate\u2019s fight, flight, or freeze defence mechanism to circumvent a snake\u2019s squeeze or bite. Numerous laboratory studies show that humans and primates both sense and visually detect snakes more rapidly than other threatening stimuli (Van Le Et al., 2013). These experiments show that snakes elicited the strongest, fastest responses (Van Le Et al., 2013). This is known as \u2018Snake Detection Theory\u2019 and is the evolution of the primate\u2019s complex brain, visual acuity, and rapid motor responses towards snakes in its environment that are the adaptations needed to live successfully as arboreal beings. It is not fortuitous then, that primates that never coexisted with venomous snakes, such as lemurs in Madagascar, have less visual acuity, better olfaction and smaller brains. Within Isbell\u2019s work, a collaborative study by a group of neuroscientists tested this hypothesis and found that, indeed, there is higher neural firing and activity in multiple areas of the primate brain, notably in the pulvinar, a region\u00a0 responsible for visual attention and oculomotor behaviour (Isbell, L., 2009).<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 509px\" class=\"wp-caption alignright\"><img src=\"https:\/\/www.worldhistory.org\/img\/r\/p\/750x750\/4941.jpg?v=1747098791\" alt=\"Apophis Defeated\" width=\"509\" height=\"309\" \/><figcaption class=\"wp-caption-text\">Tomb of Inherkau no. 359 Second chamber, South wall \"The great cat of Heliopolis\" killing the enemy of the sun, Apophis. source: https:\/\/www.worldhistory.org\/image\/4941\/apophis-defeated\/<\/figcaption><\/figure>\n<p>Today, the fear of snakes is widespread in humans, often shown through avoidance and disgust. A study in <em data-start=\"197\" data-end=\"244\">The Journal of Ethnobiology and Ethnomedicine<\/em> notes that snakes are over-hunted and excluded from conservation efforts worldwide (Cer\u00edaco, 2012). While cultural factors shape our sentiments, instinct also plays a role\u2014such as the developed avoidance behaviors toward threats like snakes. This blend of instinct and cultural influence is not only seen in behavior but also deeply embedded in the stories we tell. Many cultures depict mythological snakes as harbingers of death or chaos. In the Bible, Satan becomes a snake to tempt Eve. Norse mythology features J\u00f6rmungandr, the world serpent who signals the apocalypse. Egyptian myth tells of Apophis, who battles the sun god Ra nightly. Though sources vary, these myths consistently portray snakes as threats. As such, the widespread fear of snakes may reflect both evolutionary and cultural influences. Understood as an adaptive response inherited from primate ancestors\u2014who developed avoidance behaviors toward potentially dangerous stimuli\u2014and reinforced through myths and religious narratives, the enduring presence of snakes as potent figures of fear across human societies and primate groups highlights the complex intertwining of instinct and cultural meaning in shaping human behavior.<\/p>\n<\/div>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Early Hominins: Sahelanthropus and Orrorin<\/strong><\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">We see evidence for bipedalism in some of the earliest fossil hominins, dated from within our estimates of our divergence from chimpanzees. These hominins, however, also indicate evidence for arboreal locomotion.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The earliest dated hominin find (between 6 mya and 7 mya, based on radiometric dating of volcanic tufts) has been argued to come from Chad and is named <strong><em>Sahelanthropus tchadensis<\/em> <\/strong>(Figure 9.8; Brunet et al. 1995). The initial discovery was made in 2001 by Ahounta Djimdoumalbaye and announced in <em>Nature<\/em> in 2002 by a team led by French paleontologist Michel Brunet. The find has a small cranial capacity (360 cc) and smaller canines than those in extant great apes, though they are larger and pointier than those in humans. <span style=\"background-color: #ffff00\">This implies strongly that, over evolutionary time, the need for display and dominance among males has reduced, as has our sexual dimorphism<\/span>. A short cranial base and a foramen magnum that is more humanlike in positioning have been argued to indicate upright walking.<\/span><\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 640px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-289\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.82.jpg\" alt=\"Four views of a beige-colored skull are seen against a black background.\" width=\"640\" height=\"640\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.8: Sahelanthropus tchadensis exhibits a set of derived features, including a long, low cranium; a small, ape-sized braincase; and relatively reduced prognathism. Credit: aa <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 anterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 posterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 inferior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; and d <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 lateral left view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Initially, the inclusion of <em>Sahelanthropus<\/em> in the hominin family was debated by researchers, since the evidence for bipedalism is based on cranial evidence alone, which is not as convincing as postcranial evidence. Yet, a femur (thigh bone) and ulnae (upper arm bones) thought to belong to <em>Sahelanthropus<\/em> was discovered in 2001 (although not published until 2022). These bones may support the idea that the hominin was in fact a terrestrial biped with arboreal capabilities and behaviors (Daver et al. 2022).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Orrorin tugenensis<\/em><\/strong> (Orrorin meaning \u201coriginal man\u201d), dated to between 6 mya and 5.7 mya, was discovered near Tugen Hills in Kenya in 2000. Smaller <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1340\">cheek teeth<\/a><\/strong> (molars and premolars) than those in even more recent hominins, thick enamel, and reduced, but apelike, canines characterize this species. This is the first species that clearly indicates adaptations for bipedal locomotion, with fragmentary leg, arm, and finger bones having been found but few cranial remains. One of the most important elements discovered was a proximal femur, BAR 1002'00. The femur is the thigh bone, and the proximal part is that which articulates with the pelvis; this is very important for studying posture and locomotion. This femur indicates that <em>Ororrin<\/em> was bipedal, and recent studies suggest that it walked in a similar way to later <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1404\">Pliocene<\/a><\/strong> hominins. Some have argued that features of the finger bones suggest potential tool-making capabilities, although many researchers argue that these features are also consistent with climbing.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Early Hominins: The Genus <em>Ardipithecus<\/em><\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Another genus, <em>Ardipithecus<\/em>, is argued to be represented by at least two species: <em>Ardipithecus (Ar.) ramidus <\/em>and <em>Ar. kadabba<\/em>.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Ardipithecus ramidus<\/em><\/strong> (\u201cramid\u201d means root in the Afar language) is currently the best-known of the earliest hominins (Figure 9.9). Unlike <em>Sahelanthropus<\/em> and<em> Orrorin<\/em>, this species has a large sample size of over 110 specimens from Aramis alone. Dated to 4.4 mya, <em>Ar. ramidus<\/em> was found in Ethiopia (in the Middle Awash region and in Gona). This species was announced in 1994 by American palaeoanthropologist Tim White, based on a partial female skeleton nicknamed \u201cArdi\u201d (ARA-VP-6\/500; White et al. 1994). Ardi demonstrates a mosaic of ancestral and derived characteristics in the postcrania. For instance, she had an opposable big toe (hallux), similar to chimpanzees (i.e., more ancestral), which could have aided in climbing trees effectively. However, the pelvis and hip show that she could walk upright (i.e., it is derived), supporting her hominin status. A small brain (300 cc to 350 cc), midfacial projection, and slight prognathism show retained ancestral cranial features, but the cheek bones are less flared and robust than in later hominins.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 706px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-1310\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.9-scaled-1.jpg\" alt=\"Skull cast and partial skeleton with photographs of some bones and line drawings of others.\" width=\"706\" height=\"453\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.9a and b: Researchers believe that Ardipithecus ramidus was able to walk upright, although not as efficiently as later humans. It possessed the musculature required for tree climbing, and while moving quadrupedally, it likely placed weight on the palms of the hands rather than on the knuckles. Credit: a. <a class=\"rId61\" href=\"https:\/\/boneclones.com\/product\/ardipithecus-ramidus-skull-BH-039\">Ardipithecus ramidus Skull<\/a> by <a class=\"rId62\" href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a> is used by permission and available here under a <a class=\"rId63\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>; b. <a class=\"rId64\" href=\"https:\/\/boneclones.com\/product\/ardipithecus-ramidus-skull-BH-039\">Artist\u2019s rendition of \u201cArdi\u201d skeleton<\/a> by <a class=\"rId65\" href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a> is used by permission and available here under a <a class=\"rId66\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong><em>Ardipithecus kadabba<\/em><\/strong> (the species name means \u201coldest ancestor\u201d in the Afar language) is known from localities on the western margin of the Middle Awash region, the same locality where <em>Ar. ramidus<\/em> has been found. Specimens include mandibular fragments and isolated teeth as well as a few postcranial elements from the Asa Koma (5.5 mya to 5.77 mya) and Kuseralee Members (5.2 mya), well-dated and understood (but temporally separate) volcanic layers in East Africa. This species was discovered in 1997 by paleoanthropologist Dr. Yohannes Haile-Selassie. Originally these specimens were referred to as a subspecies of <em>Ar. ramidus<\/em>. In 2002, six teeth were discovered at Asa Koma and the dental-wear patterns confirmed that this was a distinct species, named <em>Ar. kadabba,<\/em> in 2004. One of the postcranial remains recovered included a 5.2 million-year-old toe bone that demonstrated features that are associated with toeing off (pushing off the ground with the big toe leaving last) during walking, a characteristic unique to bipedal walkers. However, the toe bone was found in the Kuseralee Member, and therefore some doubt has been cast by researchers about its association with the teeth from the Asa Koma Member.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"background-color: #ff99cc\"><strong><span style=\"color: #000000\">Bipedal Trends in Early Hominins: Summary<\/span><\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff99cc\">Trends toward bipedalism are seen in our earliest hominin finds. However, many specimens also indicate retained capabilities for climbing. Trends include a larger, more robust hallux; a more compact foot, with an arch; a robust, long femur, angled at the knee; a robust tibia; a bowl-shaped pelvis; and a more anterior foramen magnum. While the level of bipedality in <em>Salehanthropus<\/em> <em>tchadenisis<\/em> is debated since there are few fossils and no postcranial evidence, <em>Orrorin tugenensis<\/em> and <em>Ardipithecus<\/em> <em>k<\/em><em>adabba <\/em>show clear indications of some of these bipedal trends. However, some retained ancestral traits, such as an opposable hallux in <em>Ardipithecus<\/em>, indicate some retention in climbing ability.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Derived Adaptations: Early Hominin Dention<\/span><\/h2>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">The Importance of Teeth<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Teeth are abundant in the fossil record, primarily because they are already highly mineralized as they are forming, far more so than even bone. Because of this, teeth preserve readily. And, because they preserve readily, they are well-studied and better understood than many skeletal elements. In the sparse hominin (and primate) fossil record, teeth are, in some cases, all we have.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Teeth also reveal a lot about the individual from whom they came. We can tell what they evolved to eat, to which other species they may be closely related, and even, to some extent, the level of sexual dimorphism, or general variability, within a given species. This is powerful information that can be contained in a single tooth. With a little more observation, the wearing patterns on a tooth can tell us about the diet of the individual in the weeks leading up to its death. Furthermore, the way in which a tooth is formed, and the timing of formation, can reveal information about changes in diet (or even mobility) over infancy and childhood, using isotopic analyses. When it comes to our earliest hominin relatives, this information is vital for understanding how they lived.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The purpose of comparing different hominin species is to better understand the functional morphology as it applies to dentition. In this, we mean that the morphology of the teeth or masticatory system (which includes jaws) can reveal something about the way in which they were used and, therefore, the kinds of foods these hominins ate. When comparing the features of hominin groups, it is worth considering modern analogues (i.e., animals with which to compare) to make more appropriate assumptions about diet. In this way, hominin dentition is often compared with that of chimpanzees and gorillas (our close ape relatives), as well as with that of modern humans.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The most divergent group, however, is humans. Humans around the world have incredibly varied diets. Among hunter-gatherers, it can vary from a honey- and plant-rich diet, as seen in the Hadza in Tanzania, to a diet almost entirely reliant on animal fat and protein, as seen in Inuits in polar regions of the world. We are therefore considered generalists, more general than the largely <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1342\"><strong>frugivorous<\/strong><\/a> (fruit-eating) chimpanzee or the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1344\"><strong>folivorous<\/strong><\/a> (foliage-eating) gorilla, as discussed in Chapter 5.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">One way in which all humans are similar is our reliance on the processing of our food. We cut up and tear meat with tools using our hands, instead of using our front teeth (incisors and canines). We smash and grind up hard seeds, instead of crushing them with our hind teeth (molars). This means that, unlike our ape relatives, we can rely more on developing tools to navigate our complex and varied diets. <span style=\"text-decoration: underline\">(We could say)<\/span> Our brain, therefore, is our primary masticatory organ. Evolutionarily, our teeth have reduced in size and our faces are flatter, or more <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1346\">orthognathic<\/a>, <\/strong>partially in response to our increased reliance on our hands and brain to process food. Similarly, a reduction in teeth and a more generalist dental morphology could also indicate an increase in softer and more variable foods, such as the inclusion of more meat. <span style=\"background-color: #ffff00\">These trends begin early on in our evolution.<\/span> The link has been made between some of the earliest evidence for stone tool manufacture, the earliest members of our genus, and the features that we associate with these specimens.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">General Dental Trends in Early Hominins<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Several trends are visible in the dentition of early hominins. However, all tend to have the same <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1348\"><strong>dental formula<\/strong><\/a>. The dental formula tells us how many of each tooth type are present in each quadrant of the mouth. Going from the front of the mouth, this includes the square, flat <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1160\">incisors<\/a><\/strong>; the pointy <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1166\">canines<\/a><\/strong>; the small, flatter <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1350\"><strong>premolars<\/strong><\/a>; and the larger hind <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1164\">molars<\/a><\/strong>. In many primates, from Old World monkeys to great apes, the typical dental formula is 2:1:2:3. This means that if we divide the mouth into quadrants, each has two incisors, one canine, two premolars, and three molars. The eight teeth per quadrant total 32 teeth in all (although some humans have fewer teeth due to the absence of their wisdom teeth, or third molars).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 380px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image24.png\" alt=\"Anterior view of the lower face of a person showing their teeth.\" width=\"380\" height=\"253\" \/><figcaption class=\"wp-caption-text\">Figure 9.10: In humans, our canines are often a similar size to our incisors. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Adult_human_teeth.jpg\">Adult human teeth<\/a> by <a href=\"https:\/\/www.genusfotografen.se\/\">Genusfotografen<\/a> (Tomas Gunnarsson) through <a href=\"https:\/\/wikimedia.se\/\">Wikimedia Sverige<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p><span style=\"color: #000000\">The morphology of the individual teeth is where we see the most change. Among primates, large incisors are associated with food procurement or preparation (such as biting small fruits), while small incisors indicate a diet that may contain small seeds or leaves (where the preparation is primarily in the back of the mouth). Most hominins have relatively large, flat, vertically aligned incisors that <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1396\">occlude<\/a> <\/strong>(touch) relatively well, forming a \u201cbite.\u201d This differs from, for instance, the orangutan, whose teeth stick out (i.e.<em>,<\/em> are <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1352\"><strong>procumbent<\/strong><\/a>).<\/span><\/p>\n<p>&nbsp;<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">While the teeth are often aligned with diet, the canines may be misleading in that regard. We tend to associate pointy, large canines with the ripping required for meat, and the reduction (or, in some animals, the absence) of canines as indicative of herbivorous diets. In humans, our canines are often a similar size to our incisors and therefore considered <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1398\">incisiform<\/a><\/strong> (Figure 9.10). However, our closest relatives all have very long, pointy canines, particularly on their upper dentition. This is true even for the gorilla, which lives almost exclusively on plants. The canines in these instances reveal more about social structure and sexual dimorphism than diet, as large canines often signal <span style=\"background-color: #ffff00\">dominance<\/span>.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Early on in human evolution, we see a reduction in canine size. <em>Sahelanthropus tchadensis<\/em> and <em>Orrorin tugenensis<\/em> both have smaller canines than those in extant great apes, yet the canines are still larger and pointier than those in humans or more recent hominins. <span style=\"background-color: #ffff00\">This implies strongly that, over evolutionary time, the need for display and dominance among males has reduced, as has our sexual dimorphism.<\/span> In <em>Ardipithecus ramidus<\/em>, there is no obvious difference between male and female canine size, yet they are still slightly larger and pointier than in modern humans. This implies a less sexually dimorphic social structure in the earlier hominins relative to modern-day chimpanzees and gorillas.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Along with a reduction in canine size is the reduction or elimination of a canine <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1172\">diastema<\/a>:<\/strong> a gap between the teeth on the mandible that allows room for elongated teeth on the maxilla to \u201cfit\u201d in the mouth. Absence of a diastema is an excellent indication of a reduction in canine size. In animals with large canines (such as baboons), there is also often a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1354\"><strong>honing P3<\/strong><\/a>, where the first premolar (also known as P3 for evolutionary reasons) is triangular in shape, \u201csharpened\u201d by the extended canine from the upper dentition. This is also seen in some early hominins: <em>Ardipithecus<\/em>, for example, has small canines that are almost the same height as its incisors, although still larger than those in recent hominins.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The hind dentition, such as the bicuspid (two cusped) premolars or the much larger molars, are also highly indicative of a generalist diet in hominins. Among the earliest hominins, the molars are larger than we see in our genus, increasing in size to the back of the mouth and angled in such a way from the much smaller anterior dentition as to give these hominins a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1356\"><strong>parabolic<\/strong><\/a> (V-shaped) dental arch. This differs from our living relatives and some early hominins, such as <em>Sahelanthropus<\/em>, whose molars and premolars are relatively parallel between the left and right sides of the mouth, creating a U-shape.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Among more recent early hominins, the molars are larger than those in the earliest hominins and far larger than those in our own genus, <em>Homo.<\/em> Large, short molars with thick <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1358\"><strong>enamel<\/strong><\/a> allowed our early cousins to grind fibrous, coarse foods, such as sedges, which require plenty of chewing. This is further evidenced in the low <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1176\">cusps<\/a>,<\/strong> or ridges, on the teeth, which are ideal for chewing. In our genus, the hind dentition is far smaller than in these early hominins. Our teeth also have medium-size cusps, which allow for both efficient grinding and tearing\/shearing meats.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Understanding the dental morphology has allowed researchers to extrapolate very specific behaviors of early hominins. It is worth noting that while teeth preserve well and are abundant, a slew of other morphological traits additionally provide evidence for many of these hypotheses. Yet there are some traits that are ambiguous. For instance, while there are definitely high levels of sexual dimorphism in <em>Au. afarensis<\/em>, discussed in the next section, the canine teeth are reduced in size, implying that while canines may be useful indicators for sexual dimorphism, it is also worth considering other evidence.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff99cc\">In summary, trends among early hominins include a reduction in procumbency, reduced hind dentition (molars and premolars), a reduction in canine size (more incisiform with a lack of canine diastema and honing P3), flatter molar cusps, and thicker dental enamel. All early hominins have the ancestral dental formula of 2:1:2:3. These trends are all consistent with a generalist diet, incorporating more fibrous foods.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Special Topic: Contested Species<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Many named species are highly debated and argued to have specimens associated with a more variable <em>Au. afarensis <\/em>or <em>Au. anamensis<\/em> species. Sometimes these specimens are dated to times when, or found in places in which, there are \u201cgaps\u201d in the palaeoanthropological record. These are argued to represent chronospecies or variants of <em>Au. afarensis<\/em>. However, it is possible that, with more discoveries, the distinct species types will hold.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><strong><em>Australopithecus bahrelghazali<\/em><\/strong> is dated to within the time period of <em>Au. afarensi<\/em>s (3.6 mya; Brunet et al. 1995) and was the first Australopithecine to be discovered in Chad in central Africa. Researchers argue that the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1360\">holotype<\/a><\/strong>, whom discoverers have named \u201cAbel,\u201d falls under the range of variation of <em>Au. afarensis<\/em> and therefore that <em>A. bahrelghazali<\/em> does not fall into a new species (Lebatard et al. 2008). If \u201cAbel\u201d is a member of <em>Au. afarensis<\/em>, the geographic range of the species would be greatly extended.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">On a different note, <strong><em>Australopithecus <\/em><\/strong><strong><em>deyiremada<\/em><\/strong> (meaning \u201cclose relative\u201d in the Ethiopian language of Afar) is dated to 3.5 mya to 3.3 mya and is based on fossil mandible bones discovered in 2011 in Woranso-Mille (in the Afar region of Ethiopia) by Yohannes Haile-Selassie, an Ethiopian paleoanthropologist (Haile-Selassie et al. 2019). The discovery indicated, in contrast to <em>Au. afarensis<\/em>, smaller teeth with thicker enamel (potentially suggesting a harder diet) as well as a larger mandible and more projecting cheekbones. This find may be evidence that more than one closely related hominin species occupied the same region at the same temporal period (Haile-Selassie et al. 2015; Spoor 2015) or that other <em>Au. afarensis<\/em> specimens have been incorrectly designated. However, others have argued that this species has been prematurely identified and that more evidence is needed before splitting the taxa, since the variation appears subtle and may be due to slightly different niche occupations between populations over time.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><strong><em>Australopithecus garhi<\/em><\/strong> is another species found in the Middle Awash region of Ethiopia. It is currently dated to 2.5 mya (younger than <em>Au. afarensis<\/em>). Researchers have suggested it fills in a much-needed temporal \u201cgap\u201d between hominin finds in the region, with some anatomical differences, such as a relatively large cranial capacity (450 cc) and larger hind dentition than seen in other gracile Australopithecines. Similarly, the species has been argued to have longer hind limbs than <em>Au. afarensis<\/em>, although it was still able to move arboreally (Asfaw et al. 1999). However, this species is not well documented or understood and is based on only several fossil specimens. More astonishingly, crude stone tools resembling Oldowan (which will be described later) have been found in association with <em>Au. garhi<\/em>. While lacking some of the features of the Oldowan, this is one of the earliest technologies found in direct association with a hominin.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><strong><em>Kenyanthopus<\/em><\/strong><strong><em> platyops<\/em><\/strong> (the name \u201cplatyops\u201d refers to its flatter-faced appearance) is a highly contested genus\/species designation of a specimen (KNM-WT 40000) from Lake Turkana in Kenya, discovered by Maeve Leakey in 1999 (Figure 9.11). Dated to between 3.5 mya and 3.2 mya, some have suggested this specimen is an <em>Australopithecus<\/em>, perhaps even <em>Au.<\/em> <em>afarensis<\/em> (with a brain size which is difficult to determine, yet appears small), while still others have placed this specimen in <em>Homo <\/em>(small dentition and flat-orthognathic face). While taxonomic placing of this species is quite divided, the discoverers have argued that this species is ancestral to <em>Homo<\/em>, in particular to <em>Homo <\/em><em>ruldolfensis<\/em> (Leakey et al. 2001). Some researchers have additionally associated the earliest tool finds from Lomekwi, Kenya, temporally (3.3 mya) and in close geographic proximity to this specimen.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 579px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-292 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.11.jpg\" alt=\"Four views of an ancient skull are shown on a black background.\" width=\"579\" height=\"579\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.11: This specimen, KNM WT 40000 (Kenyanthopus platyops), has small detention, a small brain case, and a relatively flat face. Its genus\/species designation remains contested. Credit: a. <a class=\"rId76\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId77\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 anterior view<\/a> by \u00a9<a class=\"rId78\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId79\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId80\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId81\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId82\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 superior view<\/a> by \u00a9<a class=\"rId83\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId84\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId85\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a class=\"rId86\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId87\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 lateral left view<\/a> by \u00a9<a class=\"rId88\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId89\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId90\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; d. <a class=\"rId91\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId92\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 inferior view<\/a> by \u00a9<a class=\"rId93\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId94\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId95\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">The Genus <em>Australopithecus<\/em><br \/>\n<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The Australopithecines are a diverse group of hominins, comprising various species. <em>Australopithecus<\/em> is the given group or genus name. It stems from the Latin word <em>Australo<\/em>, meaning \u201csouthern,\u201d and the Greek word <em>pithecus,<\/em> meaning \u201cape.\u201d Within this section, we will outline these differing species\u2019 geological and temporal distributions across Africa, unique derived and\/or shared traits, and importance in the fossil record.<\/span><\/p>\n<figure style=\"width: 381px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image37-2.png\" alt=\"A skull has a pronounced sagittal crest, flaring cheekbones, and large hind teeth.\" width=\"381\" height=\"585\" \/><figcaption class=\"wp-caption-text\">Figure 9.12: Robust Australopithecines such as Paranthropus boisei had large molars and chewing muscles. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Paranthropus_boisei_skull.jpg\">Paranthropus boisei skull<\/a> by Durova is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/deed.en\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Between 3 mya and 1 mya, there seems to be differences in dietary strategy between different species of hominins designated as Australopithecines. A pattern of larger posterior dentition (even relative to the incisors and canines in the front of the mouth), thick enamel, and cranial evidence for extremely large chewing muscles is far more pronounced in a group known as the robust australopithecines. This pattern is extreme<span style=\"text-decoration: underline\">ly<\/span> relative to their earlier contemporaries or predecessors, the gracile australopithecines<strong>,<\/strong> and is certainly larger than those seen in early <em>Homo<\/em>, which emerged during this time. This pattern of incredibly large hind dentition (and very small anterior dentition) has led people to refer to robust australopithecines as <strong>megadont<\/strong> hominins (Figure 9.12).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Because of these differences, this section has been divided into \u201cgracile\u201d and \u201crobust\u201d Australopithecines, highlighting the morphological differences between the two groups (which many researchers have designated as separate genera: <em>Australopithecus<\/em> and <em>Paranthropus<\/em>, respectively) and then focusing on the individual species. It is worth noting, however, that not all researchers accept these clades as biologically or genetically distinct, with some researchers insisting that the relative gracile and robust features found in these species are due to parallel evolutionary events toward similar dietary niches.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Despite this genus\u2019 ancestral traits and small cranial capacity, all members show evidence of bipedal locomotion. It is generally accepted that <em>Australopithecus <\/em>species display varying degrees of arborealism along with bipedality.<\/span><\/p>\n<h3 class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Gracile Australopithecines<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">This section describes individual species from across Africa. These species are called \u201c<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1406\">gracile<\/a> <\/strong>australopithecines\u201d because of their smaller and less robust features compared to the divergent \u201c<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1408\"><strong>robust<\/strong><\/a>\u201d group. Numerous Australopithecine species have been named, but some are only based on a handful of fossil finds, whose designations are controversial.<\/span><\/p>\n<h4 class=\"import-Normal\"><em><span style=\"color: #000000\">East African Australopithecines<\/span><\/em><\/h4>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">East African Australopithecines are found throughout the EARS, and they include the earliest species associated with this genus. Numerous fossil-yielding sites, such as Olduvai, Turkana, and Laetoli, have excellent, datable stratigraphy, owing to the layers of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1362\"><strong>volcanic tufts<\/strong> <\/a> that have accumulated over millions of years. <span style=\"background-color: #ff9900\">These tufts may be dated using absolute dating techniques, such as Potassium-Argon dating (described in Chapter 7). This means that it is possible to know a relatively refined date for any fossil if the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1364\"><strong>context<\/strong> <\/a> (i.e., exact location) of that find is known.<\/span> Similarly, comparisons between the faunal assemblages of these stratigraphic layers have allowed researchers to chronologically identify environmental changes.<\/span><\/p>\n<figure style=\"width: 313px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image20-1-1.png\" alt=\"Occlusal view of an Au. anamensis mandible, with relatively large teeth, including canines.\" width=\"313\" height=\"313\" \/><figcaption class=\"wp-caption-text\">Figure 9.13: As seen in this mandible of KNM-KP 29281, Australopithecus anamensis had relatively large canine teeth. Credit: <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20anamensis\/KNM-KP%2029281\">Australopithecus anamensis: KNM-KP 29281 occlusal view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p><span style=\"color: #000000\">The earliest known Australopithecine is dated to 4.2 mya to 3.8 mya. <strong><em>Australopithecus anamensis<\/em><\/strong> (after \u201cAnam,\u201d meaning \u201clake\u201d from the Turkana region in Kenya; Leakey et al. 1995; Patterson and Howells 1967) is currently found from sites in the Turkana region (Kenya) and Middle Awash (Ethiopia; Figure 9.13). Recently, a 2019 find from Ethiopia, named MRD, after Miro Dora where it was found, was discovered by an Ethiopian herder named Ali Bereino. It is one of the most complete cranial finds of this species (Ward et al. 1999). A small brain size (370 cc), relatively large canines, projecting cheekbones, and earholes show more ancestral features as compared to those of more recent Australopithecines. The most important element discovered with this species is a fragment of a tibia (shinbone), which demonstrates features associated with weight transfer during bipedal walking. Similarly, the earliest found hominin femur belongs to this species. Ancestral traits in the upper limb (such as the humerus) indicate some retained arboreal locomotion.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Some researchers suggest that <em>Au. anamensis<\/em> is an intermediate form of the chronospecies that becomes <em>Au. afarensis<\/em>, evolving from <em>Ar. ramidus<\/em>. However, this is debated, with other researchers suggesting morphological similarities and affinities with more recent species instead. Almost 100 specimens, representing over 20 individuals, have been found to date (Leakey et al. 1995; McHenry 2009; Ward et al. 1999).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Au. afarensis<\/em><\/strong> is one of the oldest and most well-known australopithecine species and consists of a large number of fossil remains. <em>Au. afarensis<\/em> (which means \u201cfrom the Afar region\u201d) is dated to between 2.9 mya and 3.9 mya and is found in sites all along the EARS system, in Tanzania, Kenya, and Ethiopia (Figure 9.14). The most famous individual from this species is a partial female skeleton discovered in Hadar (Ethiopia), later nicknamed \u201cLucy,\u201d after the Beatles\u2019 song \u201cLucy in the Sky with Diamonds,\u201d which was played in celebration of the find (Johanson et al. 1978; Kimbel and Delezene 2009). This skeleton was found in 1974 by Donald Johanson and dates to approximately 3.2 mya. In addition, in 2002 a juvenile of the species was found by Zeresenay Alemseged and given the name \u201cSelam\u201d (meaning \u201cpeace,\u201d DIK 1-1), though it is popularly known as \u201cLucy\u2019s Child\u201d or as the \u201cDikika Child\u201d (Alemseged et al. 2006). Similarly, the \u201cLaetoli Footprints\u201d (discussed in Chapter 7; Hay and Leakey 1982; Leakey and Hay 1979) have drawn much attention.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 643px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-295 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.14.jpg\" alt=\"Two images of life-like reconstructions of female and male Au. afarensis.\" width=\"643\" height=\"322\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.14 a-b: Artistic reconstructions of Australopithecus afarensis by artist John Gurche. Female \u201cLucy\u201d is left and a male is on the right. Credit: a. <a class=\"rId106\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Australopithecus afarensis, \u201cLucy,\u201d adult female. Reconstruction based on AL-288-1 by artist John Gurche, front view close-up<\/a> by <a class=\"rId107\" href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?\u201d] is <a class=\"rId108\" href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>; b. <a class=\"rId109\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Australopithecus afarensis, adult male. Reconstruction based on <\/a><a class=\"rId110\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">AL444-2<\/a><a class=\"rId111\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\"> by John Gurche<\/a> by <a class=\"rId112\" href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?\u201d] is <a class=\"rId113\" href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>.<\/figcaption><\/figure>\n<figure style=\"width: 320px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image40.png\" alt=\"A partial skeleton includes bones of the cranium, mandible, and postcranium.\" width=\"320\" height=\"772\" \/><figcaption class=\"wp-caption-text\">Figure 9.15: The humanlike femoral neck, valgus knee, and bowl-shaped hip seen in the \u201cLucy\u201d skeleton indicates that Australopithecus afarensis was bipedal. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Lucy_blackbg.jpg\">Lucy blackbg<\/a> [AL 288-1, Australopithecus afarensis, cast from Museum national d'histoire naturelle, Paris] by 120 is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The canines and molars of <em>Au. afarensis<\/em> are reduced relative to great apes but are larger than those found in modern humans (indicative of a generalist diet); in addition, <em>Au. afarensis <\/em>has a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1366\"><strong>prognathic<\/strong> <\/a> face (the face below the eyes juts anteriorly) and robust facial features that indicate relatively strong chewing musculature (compared with <em>Homo<\/em>) but which are less extreme than in <em>Paranthropus<\/em>. Despite a reduction in canine size in this species, large overall size variation indicates high levels of sexual dimorphism.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Skeletal evidence indicates that this species was bipedal, as its pelvis and lower limb demonstrate a humanlike femoral neck, valgus knee, and bowl-shaped hip (Figure 9.15). <span style=\"background-color: #ccffcc\">More evidence of bipedalism is found in the footprints of this species<\/span>. <em>Au. afarensis<\/em> is associated with the Laetoli Footprints, <span style=\"background-color: #ff9900\">a 24-meter trackway of hominin fossil footprints preserved in volcanic ash discovered by Mary Leakey in Tanzania and dated to 3.5 mya to 3 mya. This set of prints is thought to have been produced by three bipedal individuals as there are no knuckle imprints, no opposable big toes, and a clear arch is present. The infants of this species are thought to have been more arboreal than the adults, as discovered through analyses of the foot bones of the Dikika Child dated to 3.32 mya (Alemseged et al. 2006).<\/span><\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Although not found in direct association with stone tools, potential evidence for cut marks on bones, found at Dikika, and dated to 3.39 mya indicates a possible temporal\/ geographic overlap between meat eating, tool use, and this species. However, this evidence is fiercely debated. Others have associated the cut marks with the earliest tool finds from Lomekwi, Kenya, temporally (3.3 mya) and in close geographic proximity to this species.<\/span><\/p>\n<h4 class=\"import-Normal\"><em><span style=\"color: #000000\">South African Australopithecines<\/span><\/em><\/h4>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Since the discovery of the Taung Child, there have been numerous Australopithecine discoveries from the region known as \u201cThe Cradle of Humankind,\u201d which was recently given UNESCO World Heritage Site status as \u201cThe Fossil Hominid Sites of South Africa.\u201d The limestone caves found in the Cradle allow for the excellent preservation of fossils. Past animals navigating the landscape and falling into cave openings, or caves used as dens by carnivores, led to the accumulation of deposits over millions of years. Many of the hominin fossils, encased in <strong>breccia<\/strong> (hard, calcareous sedimentary rock), are recently exposed from limestone quarries mined in the previous century. This means that extracting fossils requires excellent and detailed exposed work, often by a team of skilled technicians.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">While these sites have historically been difficult to date, with mixed assemblages accumulated over large time periods, advances in techniques such as uranium-series dating have allowed for greater accuracy. Historically, the excellent faunal record from East Africa has been used to compare sites based on <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1368\"><strong>relative dating<\/strong>,<\/a> whereby environmental and faunal changes and extinction events allow us to know which hominin finds are relatively younger or older than others.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The discovery of the Taung Child in 1924 (discussed in the Special Topic box \u201cThe Taung Child\u201d below) shifted the focus of palaeoanthropological research from Europe to Africa, although acceptance of this shift was slow (Broom 1947; Dart 1925). The species to which it is assigned, <strong><em>Australopithecus africanus<\/em><\/strong> (name meaning \u201cSouthern Ape of Africa\u201d), is currently dated to between 3.3 mya and 2.1 mya (Pickering and Kramers 2010), with discoveries from Sterkfontein, Taung, Makapansgat, and Gladysvale in South Africa (Figure 9.16). A relatively large brain (400 cc to 500 cc), small canines without an associated diastema, and more rounded cranium and smaller teeth than <em>Au. afarensis<\/em> indicate some derived traits. Similarly, the postcranial remains (in particular, the pelvis) indicate bipedalism. However, the sloping face and curved phalanges (indicative of retained arboreal locomotor abilities) show some ancestral features. Although not in direct association with stone tools, a 2015 study noted that the trabecular bone morphology of the hand was consistent with forceful tool manufacture and use, suggesting potential early tool abilities.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 570px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image19-1.png\" alt=\"A life-like reconstruction of the face of Australopithecus africanus, smiling in anterior view.\" width=\"570\" height=\"570\" \/><figcaption class=\"wp-caption-text\">Figure 9.16: An artistic reconstruction of Australopithecus africanus by John Gurche. Credit: <a href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Australopithecus africanus. Reconstruction based on STS 5 by John Gurche <\/a>by <a href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?] is <a href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Another famous <em>Au. africanus<\/em> skull (the skull of \u201cMrs. Ples\u201d) was previously attributed to <em>Plesianthropus transvaalensis<\/em><em>, <\/em>meaning \u201cnear human from the Transvaal,\u201d the old name for Gauteng Province, South Africa (Broom 1947, 1950). The name was shortened by contemporary journalists to \u201cPles\u201d (Figure 9.17). Due to the prevailing mores of the time, the assumed female found herself married, at least in name, and has become widely known as \u201cMrs. Ples.\u201d It was later reassigned to <em>Au. africanus<\/em> and is now argued by some to be a young male rather than an adult female cranium (Thackeray 2000, Thackeray et al. 2002).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 548px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-298 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.17.jpg\" alt=\"Four views of an ancient skull are shown on a black background.\" width=\"548\" height=\"548\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.17: The \u201cMrs. Ples\u201d brain case is small in size (like apes) but its face is less prognathic; its foramen magnum is positioned more like a modern human than an African apes. Credit: a. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 anterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 posterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 superior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; and d. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 lateral right view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In 2008, nine-year-old Matthew Berger, son of paleoanthropologist Lee Berger, noted a clavicle bone in some leftover mining breccia in the Malapa Fossil Site (South Africa). After rigorous studies, the species, <strong><em>Australopithecus sediba<\/em><\/strong> (meaning \u201cfountain\u201d or \u201cwellspring\u201d in the South African language of Sesotho), was named in 2010 (Figure 9.18; Berger et al. 2010). The first type specimen belongs to a juvenile male, Karabo (MH1), but the species is known from at least six partial skeletons, from infants through adults. These specimens are currently dated to 1.97 mya (Dirks et al. 2010). The discoverers have argued that <em>Au. sediba<\/em> shows mosaic features between <em>Au. africanus<\/em> and the genus, <em>Homo<\/em>, which potentially indicates a transitional species, although this is heavily debated. These features include a small brain size (<em>Australopithecus<\/em>-like; 420 cc to 450 cc) but gracile mandible and small teeth (<em>Homo<\/em>-like). Similarly, the postcranial skeletons are also said to have mosaic features: scientists have interpreted this mixture of traits (such as a robust ankle but evidence for an arch in the foot) as a transitional phase between a body previously adapted to arborealism (particularly in evidence from the bones of the wrist) to one that adapted to bipedal ground walking. Some researchers have argued that <em>Au. sediba<\/em> shows a modern hand morphology (shorter fingers and a longer thumb), indicating that adaptations to tool manufacture and use may be present in this species.<\/span><\/p>\n<figure style=\"width: 531px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image17-1.png\" alt=\"A beige-colored skull with no mandible on a black background has some missing teeth.\" width=\"531\" height=\"400\" \/><figcaption class=\"wp-caption-text\">Figure 9.18: Australopithecus sediba shows mosaic features between Au. africanus and Homo. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Australopithecus_sediba.JPG\">Australopithecus sediba<\/a>, photo by Brett Eloff courtesy <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Profberger\">Profberger<\/a> and <a href=\"https:\/\/en.wikipedia.org\/wiki\/University_of_the_Witwatersrand\">Wits University<\/a>, is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Another famous Australopithecine find from South Africa is that of the nearly complete skeleton now known as \u201cLittle Foot\u201d (Clarke 1998, 2013). Little Foot (StW 573) is potentially the earliest dated South African hominin fossil, dating to 3.7 mya, based on radiostopic techniques, although some argue that it is younger than 3 mya (Pickering and Kramers 2010). The name is jokingly in contrast to the cryptid species \u201cbigfoot\u201d and is named because the initial discovery of four ankle bones indicated bipedality. Little Foot was discovered by Ron Clarke in 1994, when he came across the ankle bones while sorting through monkey fossils in the University of Witwatersrand collections (Clarke and Tobias 1995). He asked Stephen Motsumi and Nkwane Molefe to identify the known records of the fossils, which allowed them to find the rest of the specimen within just days of searching the Sterkfontein Caves\u2019 Silberberg Grotto.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The discoverers of Little Foot insist that other fossil finds, previously identified as <em>Au. Africanus<\/em>, be placed in this new species based on shared ancestral traits with older East African Australopithecines (Clarke and Kuman 2019). These include features such as a relatively large brain size (408 cc), robust zygomatic arch, and a flatter midface. Furthermore, the discoverers have argued that the heavy anterior dental wear patterns, relatively large anterior dentition, and smaller hind dentition of this specimen more closely resemble that of <em>Au. anamensis<\/em> or <em>Au. afarensis<\/em>. It has thus been placed in the species <strong><em>Australopithecus prometheus<\/em><\/strong>. This species name refers to a previously defunct taxon named by Raymond Dart. The species designation was, through analyzing Little Foot, revived by Ron Clarke, who insists that many other fossil hominin specimens have prematurely been placed into <em>Au. africanus<\/em>. Others say that it is more likely that <em>Au. africanus<\/em> is a more variable species and not representative of two distinct species.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\"><em>Paranthropus<\/em> \u201cRobust\u201d Australopithecines<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In the robust australopithecines, the specialized nature of the teeth and masticatory system, such as flaring zygomatic arches (cheekbones), accommodate very large temporalis (chewing) muscles. These features also include a large, broad, dish-shaped face and and a large mandible with extremely large posterior dentition (referred to as megadonts) and hyper-thick enamel (Kimbel 2015; Lee-Thorp 2011; Wood 2010). Research has revolved around the shared adaptations of these \u201crobust\u201d australopithecines, linking their morphologies to a diet of hard and\/or tough foods (Brain 1967; Rak 1988). Some argued that the diet of the robust australopithecines was so specific that any change in environment would have accelerated their extinction. The generalist nature of the teeth of the gracile australopithecines, and of early <em>Homo<\/em>, would have made them more capable of adapting to environmental change. However, some have suggested that the features of the robust australopithecines might have developed as an effective response to what are known as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1370\"><strong>fallback <\/strong><strong>foods<\/strong><\/a> in hard times rather than indicating a lack of adaptability.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are currently three widely accepted robust australopithecus or, <em>Paranthropus<\/em>, species: <em>P. aethiopic<\/em><em>us<\/em>, which has more ancestral traits, and <em>P. boisei and P. robustus<\/em>, which are more derived in their features (Strait et al. 1997; Wood and Schroer 2017). These three species have been grouped together by a majority of scholars as a single genus as they share more derived features (are more closely related to each other; or, in other words, are <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1372\"><strong>monophyletic<\/strong><\/a>) than the other australopithecines (Grine 1988; Hlazo 2015; Strait et al. 1997; Wood 2010 ). While researchers have mostly agreed to use the umbrella term <em>Paranthropus<\/em>, there are those who disagree (Constantino and Wood 2004, 2007; Wood 2010).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">As a collective, this genus spans 2.7 mya to 1.0 mya, although the dates of the individual species differ. The earliest of the Paranthropus species, <strong><em>Paranthropus aethiopicus<\/em><\/strong>, is dated to between 2.7 mya and 2.3 mya and currently found in Tanzania, Kenya, and Ethiopia in the EARS system (Figure 9.19; Constantino and Wood 2007; Hlazo 2015; Kimbel 2015; Walker et al. 1986; White 1988). It is well known because of one specimen known as the \u201cBlack Skull\u201d (KNM\u2013WT 17000), so called because of the mineral manganese that stained it black during fossilization (Kimbel 2015). As with all robust Australopithecines, <em>P. aethiopicus<\/em> has the shared derived traits of large, flat premolars and molars; large, flaring zygomatic arches for accommodating large chewing muscles (the temporalis muscle); a sagittal crest (ridge on the top of the skull) for increased muscle attachment of the chewing muscles to the skull; and a robust mandible and supraorbital torus (brow ridge). However, only a few teeth have been found. A proximal tibia indicates bipedality and similar body size to <em>Au. afarensis<\/em>. In recent years, researchers have discovered and assigned a proximal tibia and juvenile cranium (L.338y-6) to the species (Wood and Boyle 2016).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 666px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-300 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.19.jpg\" alt=\"Five views of a beige partial skull on a black background.\" width=\"666\" height=\"444\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.19: The \u201cBlack Skull\u201d (Paranthropus aethiopicus) had a large sagittal crest and large, flared zygomatic arches that indicate it had large chewing muscles and a powerful biting force. Credit: a. <a class=\"rId156\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId157\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 anterior view<\/a> by \u00a9<a class=\"rId158\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId159\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId160\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId161\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId162\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 lateral right view<\/a> by \u00a9<a class=\"rId163\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId164\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId165\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a class=\"rId166\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId167\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 superior view<\/a> by \u00a9<a class=\"rId168\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId169\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId170\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; d. <a class=\"rId171\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId172\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 posterior view<\/a> by \u00a9<a class=\"rId173\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId174\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId175\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; e. <a class=\"rId176\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId177\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 inferior view<\/a> by \u00a9<a class=\"rId178\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId179\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId180\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">First attributed as <em>Zinjanthropus boisei<\/em> (with the first discovery going by the nickname \u201cZinj\u201d or sometimes \u201cNutcracker Man\u201d), <strong><em>Paranthropus boisei<\/em><\/strong> was discovered in 1959 by Mary Leakey (see Figure 9.20 and 9.21; Hay 1990; Leakey 1959). This \u201crobust\u201d australopith species is distributed across countries in East Africa at sites such as Kenya (Koobi Fora, West Turkana, and Chesowanja), Malawi (Malema-Chiwondo), Tanzania (Olduvai Gorge and Peninj), and Ethiopia (Omo River Basin and Konso). The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1374\">hypodigm<\/a><\/strong>, sample of fossils whose features define the group, has been found by researchers to date to roughly 2.4 mya to 1.4 mya. Due to the nature of its exaggerated, larger, and more robust features, <em>P. boisei <\/em>has been termed <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1376\"><strong>hyper-robust<\/strong><\/a>\u2014that is, even more heavily built than other robust species, with very large, flat posterior dentition (Kimbel 2015). Tools dated to 2.5 mya in Ethiopia have been argued to possibly belong to this species. Despite the cranial features of <em>P. boisei<\/em> indicating a tough diet of tubers, nuts, and seeds, isotopes indicate a diet high in C4 foods (e.g., grasses, such as sedges). Another famous specimen from this species is the Peninj mandible from Tanzania, found in 1964 by Kimoya Kimeu.<\/span><\/p>\n<figure style=\"width: 557px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image14-1.png\" alt=\"Life-like reconstruction of Paranthropus boisei.\" width=\"557\" height=\"557\" \/><figcaption class=\"wp-caption-text\">Figure 9.20: Artistic reconstruction of a Paranthropus boisei, male, by John Gurche. Credit: <a href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Paranthropus boisei, male. Reconstruction based on OH 5 and KNM-ER 406 by John Gurche<\/a> by <a href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?\u201d] is <a href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>.<\/figcaption><\/figure>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 565px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-302 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.21.jpg\" alt=\"Three views of an ancient skull are shown on a black background.\" width=\"565\" height=\"565\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.21: \u201cNutcracker Man\u201d (Paranthropus boisei) had hyper-robust features including very large dentition, flaring zygomatic arches, a broad concave face. It had a powerful and extremely efficient chewing force. Credit: <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20boisei\/OH%205\">Paranthropus boisei: OH 5 anterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20boisei\/OH%205\">Paranthropus boisei: OH 5 inferior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20boisei\/OH%205\">Paranthropus boisei: OH 5 posterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Paranthropus robustus<\/em><\/strong> was the first taxon to be discovered within the genus in Kromdraai B by a schoolboy named Gert Terblanche; subsequent fossil discoveries were made by researcher Robert Broom in 1938 (Figure 9.22; Broom 1938a, 1938b, 1950), with the holotype specimen TM 1517 (Broom 1938a, 1938b, 1950; Hlazo 2018). <em>Paranthropus robustus<\/em> dates approximately from 2.0 mya to 1 mya and is the only taxon from the genus to be discovered in South Africa. Several of these fossils are fragmentary in nature, distorted, and not well preserved because they have been recovered from quarry breccia using explosives. <em>P. robustus<\/em> features are neither as \u201chyper-robust\u201d as <em>P. boisei<\/em> nor as ancestral as <em>P. aethiopicus<\/em>; instead, they have been described as being less derived, more general features that are shared with both East African species (e.g., the sagittal crest and zygomatic flaring; Rak 1983; Walker and Leakey 1988). Enamel hypoplasia is also common in this species, possibly because of instability in the development of large, thick-enameled dentition.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 572px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-303 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.22.jpg\" alt=\"Four views of a beige-colored skull are shown on a black background.\" width=\"572\" height=\"619\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.22: SK 48, a Paranthropus robustus specimen, had less derived, more general features that were not as robust as P. boisei and not as ancestral as P. aethiopicus. Credit: a. <a class=\"rId208\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId209\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 anterior view<\/a> by \u00a9<a class=\"rId210\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId211\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId212\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId213\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId214\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 superior view<\/a> by \u00a9<a class=\"rId215\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId216\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId217\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a class=\"rId218\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId219\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 inferior view<\/a> by \u00a9<a class=\"rId220\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId221\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId222\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; d. <a class=\"rId223\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId224\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 lateral left view<\/a> by \u00a9<a class=\"rId225\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId226\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId227\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Comparisons between Gracile and Robust Australopiths<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Comparisons between gracile and robust australopithecines may indicate different phylogenetic groupings or parallel evolution in several species. In general, the robust australopithecines have large temporalis (chewing) muscles, as indicated by flaring zygomatic arches, sagittal crests, and robust mandibles (jawbones). Their hind dentition is large (megadont), with low cusps and thick enamel. Within the gracile australopithecines, researchers have debated the relatedness of the species, or even whether these species should be lumped together to represent more variable or polytypic species. Often researchers will attempt to draw chronospecific trajectories, with one taxon said to evolve into another over time.<\/span><\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Special Topic: The Taung Child<\/span><\/h2>\n<figure id=\"attachment_304\" aria-describedby=\"caption-attachment-304\" style=\"width: 570px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-304 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.23.jpg\" alt=\"An ancient skull in anterior and lateral views. One view shows an imprint of the brain.\" width=\"570\" height=\"285\" \/><figcaption id=\"caption-attachment-304\" class=\"wp-caption-text\">Figure 9.23: The Taung Child has a nearly complete face, mandible, and partial endocranial cast. Credit: a. <em>A<\/em><a class=\"rId230\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\"><em>ustralopithecus africanus<\/em><\/a><a class=\"rId231\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\">: Taung 1 anterior view<\/a> by \u00a9<a class=\"rId232\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId233\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId234\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId235\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\"><em>australopithecus africanus<\/em><\/a><a class=\"rId236\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\">: Taung 1 lateral right view<\/a> by \u00a9<a class=\"rId237\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId238\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId239\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">The well-known fossil of a juvenile <em>Australopithecine<\/em>, the \u201cTaung Child,\u201d was the first early hominin evidence ever discovered and was the first to demonstrate our common human heritage in Africa (Figure 9.23; Dart 1925). The tiny facial skeleton and natural endocast were discovered in 1924 by a local quarryman in the North West Province in South Africa and were painstakingly removed from the surrounding cement-like breccia by Raymond Dart using his wife\u2019s knitting needles. When first shared with the scientific community in 1925, it was discounted as being nothing more than a young monkey of some kind. Prevailing biases of the time made it too difficult to contemplate that this small-brained hominin could have anything to do with our own history. The fact that it was discovered in Africa simply served to strengthen this bias.<\/span><\/p>\n<\/div>\n<p>&nbsp;<\/p>\n<h2><span style=\"color: #000000\">Early Tool Use and Technology<br \/>\n<\/span><\/h2>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Early Stone Age Technology (ESA)<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1378\">Early Stone Age (ESA)<\/a><\/strong> marks the beginning of recognizable technology made by our human ancestors. Stone-tool (or <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1380\"><strong>lithic<\/strong><\/a>) technology is defined by the fracturing of rocks and the manufacture of tools through a process called <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1382\"> <strong>knapping<\/strong><\/a>. The Stone Age lasted for more than 3 million years and is broken up into chronological periods called the Early (ESA), Middle (MSA), and Later Stone Ages (LSA). Each period is further broken up into a different <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1384\">techno-complex<\/a><\/strong>, a term encompassing multiple <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1386\">assemblages<\/a><\/strong> (collections of artifacts) that share similar traits in terms of artifact production and morphology. The ESA spanned the largest technological time period of human innovation from over 3 million years ago to around 300,000 years ago and is associated almost entirely with hominin species prior to modern <em>Homo sapiens. <\/em>As the ESA advanced, stone tool makers (known as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1390\"><strong>knappers<\/strong><\/a>) began to change the ways they detached <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1388\">flakes<\/a><\/strong> and eventually were able to shape artifacts into functional tools. These advances in technology go together with the developments in human evolution and cognition, dispersal of populations across the African continent and the world, and climatic changes.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In order to understand the ESA, it is important to consider that not all assemblages are exactly the same within each techno-complex: one can have multiple phases and traditions at different sites (Lombard et al. 2012). However, there is an overarching commonality between them. Within stone tool assemblages, both flakes or <strong>cores<\/strong> (the rocks from which flakes are removed) are used as tools.<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1392\"> <strong>Large Cutting Tools (LCTs)<\/strong><\/a> are tools that are shaped to have functional edges. It is important to note that the information presented here is a small fraction of what is known about the ESA, and there are ongoing debates and discoveries within archaeology.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Currently, the oldest-known stone tools, which form the techno-complex the Lomekwian, date to 3.3 mya (Harmand et al. 2015; Toth 1985). They were found at a site called Lomekwi 3 in Kenya. This techno-complex is the most recently defined and pushed back the oldest-known date for lithic technology. There is only one known site thus far and, due to the age of the site, it is associated with species prior to <em>Homo<\/em>, such as <em>Kenyanthropus platyops.<\/em> Flakes were produced through indirect percussion, whereby the knappers held a rock and hit it against another rock resting on the ground. The pieces are very chunky and do not display the same fracture patterns seen in later techno-complexes. Lomekwian knappers likely aimed to get a sharp-edged piece on a flake, which would have been functional, although the specific function is currently unknown.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Stone tool use, however, is not only understood through the direct discovery of the tools. Cut marks on fossilized animal bones may illuminate the functionality of stone tools. In one controversial study in 2010, researchers argued that cut marks on a pair of animal bones from Dikika (Ethiopia), dated to 3.4 mya, were from stone tools. The discoverers suggested that they be more securely associated, temporally, with <em>Au. afarensis<\/em>. However, others have noted that these marks are consistent with teeth marks from crocodiles and other carnivores.<\/span><\/p>\n<figure style=\"width: 324px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image29-1.png\" alt=\"A technical line drawing of an Oldowan chopper.\" width=\"324\" height=\"275\" \/><figcaption class=\"wp-caption-text\">Figure 9.24: Some scholars believe that some genera explored in this chapter were capable of producing more complex stone tools (Oldowan). Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Olduwan_Industry_Chopper_2.jpeg\">Olduwan Industry Chopper 2<\/a> by Emmyanne29 is under a <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/legalcode\">CC0 1.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The Oldowan techno-complex is far more established in the scientific literature (Leakey 1971). It is called the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1394\">Oldowan<\/a><\/strong> because it was originally discovered in Olduvai Gorge, Tanzania, but the oldest assemblage is from Gona in Ethiopia, dated to 2.6 mya (Semaw 2000). The techno-complex is defined as a core and flake industry. Like the Lomekwian, there was an aim to get sharp-edged flakes, but this was achieved through a different production method. Knappers were able to actively hold or manipulate the core being knapped, which they could directly hit using a hammerstone. This technique is known as free-hand percussion, and it demonstrates an understanding of fracture mechanics. It has long been argued that the Oldowan hominins were skillful in tool manufacture.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Because Oldowan knapping requires skill, earlier researchers have attributed these tools to members of our genus, <em>Homo<\/em>. However, some have argued that these tools are in more direct association with hominins in the genera described in this chapter (Figure 9.24).<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Invisible Tool Manufacture and Use<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The vast majority of our understanding of these early hominins comes from fossils and reconstructed paleoenvironments. It is only from 3 mya when we can start \u201clooking into their minds\u201d and lifestyles by analyzing their manufacture and use of stone tools. However, the vast majority of tool use in primates (and, one can argue, in humans) is not with durable materials like stone. All of our extant great ape relatives have been observed using sticks, leaves, and other materials for some secondary purpose (to wade across rivers, to \u201cfish\u201d for termites, or to absorb water for drinking). It is possible that the majority of early hominin tool use and manufacture may be invisible to us because of this preservation bias.<\/span><\/p>\n<h2 class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000;background-color: #ff00ff\">Chapter Summary<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000;background-color: #ff00ff\">The fossil record of our earliest hominin relatives has allowed paleoanthropologists to unpack some of the mysteries of our evolution. We now know that traits associated with bipedalism evolved before other \u201chuman-like\u201d traits, even though the first hominins were still very capable of arboreal locomotion. We also know that, for much of this time, hominin taxa were diverse in the way they looked and what they ate, and they were widely distributed across the African continent. And we know that the environments in which these hominins lived underwent many changes over this time during several warming and cooling phases.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;background-color: #ff00ff\">Yet this knowledge has opened up many new mysteries. We still need to better differentiate some taxa. In addition, there are ongoing debates about why certain traits evolved and what they meant for the extinction of some of our relatives (like the robust australopiths). The capabilities of these early hominins with respect to tool use and manufacture is also still uncertain.<\/span><\/p>\n<h2 class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Hominin Species Summaries<br \/>\n<\/span><\/h2>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Sahelanthropus tchadensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">7 mya to 6 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Chad<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The initial discovery, made in 2001.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">360 cc average<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Smaller than in extant great apes; larger and pointier than in humans. Canines worn at the tips.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A short cranial base and a foramen magnum (hole in which the spinal cord enters the cranium) that is more humanlike in positioning; has been argued to indicate upright walking.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Currently little published postcranial material.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The extent to which this hominin was bipedal is currently heavily debated. If so, it would indicate an arboreal bipedal ancestor of hominins, not a knuckle-walker like chimpanzees.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Orrorin tugenensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">6 mya to 5.7 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Tugen Hills (Kenya)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Original discovery in 2000.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Smaller cheek teeth (molars and premolars) than even more recent hominins (i.e., derived), thick enamel, and reduced, but apelike, canines.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Not many found<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Fragmentary leg, arm, and finger bones have been found. Indicates bipedal locomotion.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Potential toolmaking capability based on hand morphology, but nothing found directly.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">This is the earliest species that clearly indicates adaptations for bipedal locomotion.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Ardipithecus kadabba<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">5.2 mya to 5.8 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Middle Awash (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Discovered by Yohannes Haile-Selassie in 1997.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Larger hind dentition than in modern chimpanzees. Thick enamel and larger canines than in later hominins.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A large hallux (big toe) bone indicates a bipedal \u201cpush off.\u201d<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Faunal evidence indicates a mixed grassland\/woodland environment.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Ardipithecus ramidus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">4.4 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Middle Awash region and Gona (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">A partial female skeleton nicknamed \u201cArdi\u201d (ARA-VP-6\/500) (found in 1994).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">300 cc to 350 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Little differences between the canines of males and females (small sexual dimorphism).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Midfacial projection, slightly prognathic. Cheekbones less flared and robust than in later hominins.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Ardi demonstrates a mosaic of ancestral and derived characteristics in the postcrania. For instance, an opposable big toe similar to chimpanzees (i.e., more ancestral), which could have aided in climbing trees effectively. However, the pelvis and hip show that she could walk upright (i.e., it is derived), supporting her hominin status.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">None directly associated<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">Over 110 specimens from Aramis<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus anamensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">4.2 mya to 3.8 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Turkana region (Kenya); Middle Awash (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A 2019 find from Ethiopia, named MRD.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">370 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Relatively large canines compared with more recent Australopithecines.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Projecting cheekbones and ancestral earholes.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lower limb bones (tibia and femur) indicate bipedality; arboreal features in upper limb bones (humerus) found.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Almost 100 specimens, representing over 20 individuals, have been found to date.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus afarensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">3.9 mya to 2.9 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Afar Region, Omo, Maka, Fejej, and Belohdelie (Ethiopia); Laetoli (Tanzania); Koobi Fora (Kenya)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lucy (discovery: 1974), Selam (Dikika Child, discovery: 2000), Laetoli Footprints (discovery: 1976).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">380 cc to 430 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Reduced canines and molars relative to great apes but larger than in modern humans.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Prognathic face, facial features indicate relatively strong chewing musculature (compared with <em>Homo<\/em>) but less extreme than in <em>Paranthropus<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clear evidence for bipedalism from lower limb postcranial bones. Laetoli Footprints indicate humanlike walking. Dikika Child bones indicate retained ancestral arboreal traits in the postcrania.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">None directly, but close in age and proximity to controversial cut marks at Dikika and early tools in Lomekwi.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Au. afarensis<\/em> is one of the oldest and most well-known australopithecine species and consists of a large number of fossil remains.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus bahrelghazali<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">3.6 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Chad<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">\u201cAbel,\u201d the holotype (discovery: 1995).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Arguably within range of variation of <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus prometheus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">3.7 mya (debated)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">Sterkfontein (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">\u201cLittle Foot\u201d (StW 573) (discovery: 1994)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">408 cc (Little Foot estimate)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">Heavy anterior dental wear patterns, relatively large anterior dentition and smaller hind dentition, similar to <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">Relatively larger brain size, robust zygomatic arch, and a flatter midface.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">The initial discovery of four ankle bones indicated bipedality.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">Highly debated new species designation.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus <\/em><em>deyiremada<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">3.5 mya to 3.3 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Woranso-Mille (Afar region, Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">First fossil mandible bones were discovered in 2011 in the Afar region of Ethiopia by Yohannes Haile-Selassie.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Smaller teeth with thicker enamel than seen in <em>Au. afarensis<\/em>, with a potentially hardier diet.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Larger mandible and more projecting cheekbones than in <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Contested species designation; arguably a member of <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Kenyanthopus<\/em><em> platyops<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">3.5 mya to 3.2 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Lake Turkana (Kenya)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">KNM\u2013WT 40000 (discovered 1999)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Difficult to determine but appears within the range of <em>Australopithecus afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Small molars\/dentition (<em>Homo<\/em>-like characteristic)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Flatter (i.e., orthognathic) face<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Some have associated the earliest tool finds from Lomekwi, Kenya, temporally (3.3 mya) and in close geographic proximity to this species\/specimen.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Taxonomic placing of this species is quite divided. The discoverers have argued that this species is ancestral to <em>Homo<\/em>, in particular to <em>Homo <\/em><em>ruldolfensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Australopithecus africanus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">3.3 mya to 2.1 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Sterkfontein, Taung, Makapansgat, Gladysvale (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Taung Child (discovery in 1994), \u201cMrs. Ples\u201d (discover in 1947), Little Foot (arguable; discovery in 1994).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">400 cc to 500 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Smaller teeth (derived) relative to <em>Au. afarensis<\/em>. Small canines with no diastema.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">A rounder skull compared with <em>Au. afarensis<\/em> in East Africa. A sloping face (ancestral).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Similar postcranial evidence for bipedal locomotion (derived pelvis) with retained arboreal locomotion, e.g., curved phalanges (fingers), as seen in <em>Au. afarensis.<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">None with direct evidence.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">A 2015 study noted that the trabecular bone morphology of the hand was consistent with forceful tool manufacture and use, suggesting potential early tool abilities.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus garhi<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.5 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Middle Awash (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">450 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Larger hind dentition than seen in other gracile Australopithecines.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A femur of a fragmentary partial skeleton, argued to belong to <em>Au. garhi<\/em>, indicates this species may be longer-limbed than <em>Au. afarensis<\/em>, although still able to move arboreally.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Crude stone tools resembling Oldowan (described later) have been found in association with <em>Au. garhi<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">This species is not well documented or understood and is based on only a few fossil specimens.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Paranthropus aethiopicus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.7 mya to 2.3 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">West Turkana (Kenya); Laetoli (Tanzania); Omo River Basin (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The \u201cBlack Skull\u201d (KNM\u2013WT 17000) (discovery 1985).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain Size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">410 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>P. aethiopicus<\/em> has the shared derived traits of large flat premolars and molars, although few teeth have been found.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Large flaring zygomatic arches for accommodating large chewing muscles (the temporalis muscle), a sagittal crest for increased muscle attachment of the chewing muscles to the skull, and a robust mandible and supraorbital torus (brow ridge).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A proximal tibia indicates bipedality and similar size to <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The \u201cBlack Skull\u201d is so called because of the mineral manganese that stained it black during fossilization.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Paranthropus boisei<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.4 mya to 1.4 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Koobi Fora, West Turkana, and Chesowanja (Kenya); Malema-Chiwondo (Malawi), Olduvai Gorge and Peninj (Tanzania); and Omo River basin and Konso (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">\u201cZinj,\u201d or sometimes \u201cNutcracker Man\u201d (OH5), in 1959 by Mary Leakey. The Peninj mandible from Tanzania, found in 1964 by Kimoya Kimeu.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">500 cc to 550 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Very large, flat posterior dentition (largest of all hominins currently known). Much smaller anterior dentition. Very thick dental enamel.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Indications of very large chewing muscles (e.g., flaring zygomatic arches and a large sagittal crest).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Evidence for high variability and sexual dimorphism, with estimates of males at 1.37 meters tall and females at 1.24 meters.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Richard Leakey and Bernard Wood have both suggested that<em> P. boisei<\/em> could have made and used stone tools. Tools dated to 2.5 mya in Ethiopia have been argued to possibly belong to this species.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Despite the cranial features of <em>P. boisei<\/em> indicating a tough diet of tubers, nuts, and seeds, isotopes indicate a diet high in C4 foods (e.g., grasses, such as sedges). This differs from what is seen in<em> P. robustus<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Australopithecus sediba<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">1.97 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Malapa Fossil Site (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Karabo (MH1) (discovery in 2008)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">420 cc to 450 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Small dentition with Australopithecine cusp-spacing.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Small brain size (<em>Australopithecus<\/em>-like) but gracile mandible (<em>Homo<\/em>-like).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Scientists have interpreted this mixture of traits (such as a robust ankle but evidence for an arch in the foot) as a transitional phase between a body previously adapted to arborealism (tree climbing, particularly in evidence from the bones of the wrist) to one that adapted to bipedal ground walking.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">None of direct association, but some have argued that a modern hand morphology (shorter fingers and a longer thumb) means that adaptations to tool manufacture and use may be present in this species.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">It was first discovered through a clavicle bone in 2008 by nine-year-old Matthew Berger, son of paleoanthropologist Lee Berger.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Paranthropus robustus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.3 mya to 1 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kromdraai B, Swartkrans, Gondolin, Drimolen, and Coopers Cave (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">SK48 (original skull)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">410 cc to 530 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Large posterior teeth with thick enamel, consistent with other Robust Australopithecines. Enamel hypoplasia is also common in this species, possibly because of instability in the development of large, thick enameled dentition.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>P. robustus<\/em> features are neither as \u201chyper-robust\u201d as <em>P. boisei<\/em> or as ancestral in features as <em>P. aethiopicus<\/em>. They have been described as less derived, more general features that are shared with both East African species (e.g., the sagittal crest and zygomatic flaring).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Reconstructions indicate sexual dimorphism.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Several of these fossils are fragmentary in nature, distorted, and not well preserved, because they have been recovered from quarry breccia using explosives.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\"><strong><span style=\"color: #000000\">Review Questions<br \/>\n<\/span><\/strong><\/h2>\n<ul>\n<li class=\"import-Normal\"><span style=\"color: #000000\">What is the difference between a \u201cderived\u201d versus an \u201cancestral\u201d trait? Give an example of both, seen in <em>Au. afarensis<\/em>.<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">Which of the paleoenvironment hypotheses have been used to describe early hominin diversity, and which have been used to describe bipedalism?<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">Which anatomical features for bipedalism do we see in early hominins?<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">Describe the dentition of gracile and robust australopithecines. What might these tell us about their diets?<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">List the hominin species argued to be associated with stone tool technologies. Are you convinced of these associations? Why\/why not?<\/span><\/li>\n<\/ul>\n<\/div>\n<h2><span style=\"color: #000000\">Key Terms<\/span><\/h2>\n<p><span style=\"color: #000000\"><strong>Arboreal:<\/strong> Related to trees or woodland.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Aridification:<\/strong> Becoming increasingly arid or dry, as related to the climate or environment.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Aridity Hypothesis:<\/strong> The hypothesis that long-term aridification and expansion of savannah biomes were drivers in diversification in early hominin evolution.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Assemblage:<\/strong> A collection demonstrating a pattern. Often pertaining to a site or region.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Bipedalism:<\/strong> The locomotor ability to walk on two legs.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Breccia:<\/strong> Hard, calcareous sedimentary rock.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Canines:<\/strong> The pointy teeth just next to the incisors, in the front of the mouth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cheek teeth:<\/strong> Or hind dentition (molars and premolars).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Chronospecies:<\/strong> Species that are said to evolve into another species, in a linear fashion, over time.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Clade:<\/strong> A group of species or taxa with a shared common ancestor.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cladistics:<\/strong> The field of grouping organisms into those with shared ancestry.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Context:<\/strong> As pertaining to palaeoanthropology, this term refers to the place where an artifact or fossil is found.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cores:<\/strong> The remains of a rock that has been flaked or knapped.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cusps:<\/strong> The ridges or \u201cbumps\u201d on the teeth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Dental formula:<\/strong> A technique to describe the number of incisors, canines, premolars, and molars in each quadrant of the mouth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Derived traits:<\/strong> Newly evolved traits that differ from those seen in the ancestor.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Diastema:<\/strong> A tooth gap between the incisors and canines.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Early Stone Age (ESA):<\/strong> The earliest-described archaeological period in which we start seeing stone-tool technology.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>East African Rift System (EARS):<\/strong> This term is often used to refer to the Rift Valley, expanding from Malawi to Ethiopia. This active geological structure is responsible for much of the visibility of the paleoanthropological record in East Africa.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Enamel:<\/strong> The highly mineralized outer layer of the tooth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Encephalization:<\/strong> Expansion of the brain.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Extant:<\/strong> Currently living\u2014i.e., not extinct.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Fallback foods:<\/strong> Foods that may not be preferred by an animal (e.g., foods that are not nutritionally dense) but that are essential for survival in times of stress or scarcity.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Fauna:<\/strong> The animals of a particular region, habitat, or geological period.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Faunal assemblages:<\/strong> Collections of fossils of the animals found at a site.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Faunal turnover:<\/strong> The rate at which species go extinct and are replaced with new species.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Flake:<\/strong> The piece knocked off of a stone core during the manufacture of a tool, which may be used as a stone tool.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Flora:<\/strong> The plants of a particular region, habitat, or geological period.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Folivorous:<\/strong> Foliage-eating.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Foramen magnum:<\/strong> The large hole (foramen) at the base of the cranium, through which the spinal cord enters the skull.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Fossil:<\/strong> The remains or impression of an organism from the past.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Frugivorous:<\/strong> Fruit-eating.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Generalist:<\/strong> A species that can thrive in a wide variety of habitats and can have a varied diet.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Glacial:<\/strong> Colder, drier periods during an ice age when there is more ice trapped at the poles.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Gracile:<\/strong> Slender, less rugged, or pronounced features.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hallux:<\/strong> The big toe.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Holotype:<\/strong> A single specimen from which a species or taxon is described or named.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hominin:<\/strong> A primate category that includes humans and our fossil relatives since our divergence from extant great apes.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Honing P3:<\/strong> The mandibular premolar alongside the canine (in primates, the P3), which is angled to give space for (and sharpen) the upper canines.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hyper-robust:<\/strong> Even more robust than considered normal in the Paranthropus genus.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hypodigm:<\/strong> A sample (here, fossil) from which researchers extrapolate features of a population.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Incisiform:<\/strong> An adjective referring to a canine that appears more incisor-like in morphology.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Incisors:<\/strong> The teeth in the front of the mouth, used to bite off food.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Interglacial:<\/strong> A period of milder climate in between two glacial periods.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Isotopes:<\/strong> Two or more forms of the same element that contain equal numbers of protons but different numbers of neutrons, giving them the same chemical properties but different atomic masses.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Knappers:<\/strong> The people who fractured rocks in order to manufacture tools.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Knapping:<\/strong> The fracturing of rocks for the manufacture of tools.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Large Cutting Tool (LCT):<\/strong> A tool that is shaped to have functional edges.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Last Common Ancestor (LCA):<\/strong> The hypothetical final ancestor (or ancestral population) of two or more taxa before their divergence.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Lithic:<\/strong> Relating to stone (here to stone tools).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Lumbar lordosis:<\/strong> The inward curving of the lower (lumbar) parts of the spine. The lower curve in the human S-shaped spine.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Lumpers:<\/strong> Researchers who prefer to lump variable specimens into a single species or taxon and who feel high levels of variation is biologically real.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Megadont:<\/strong> An organism with extremely large dentition compared with body size.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Metacarpals:<\/strong> The long bones of the hand that connect to the phalanges (finger bones).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Molars:<\/strong> The largest, most posterior of the hind dentition.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Monophyletic:<\/strong> A taxon or group of taxa descended from a common ancestor that is not shared with another taxon or group.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Morphology:<\/strong> The study of the form or size and shape of things; in this case, skeletal parts.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Mosaic evolution:<\/strong> The concept that evolutionary change does not occur homogeneously throughout the body in organisms.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Obligate bipedalism:<\/strong> Where the primary form of locomotion for an organism is bipedal.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Occlude:<\/strong> When the teeth from the maxilla come into contact with the teeth in the mandible.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Oldowan:<\/strong> Lower Paleolithic, the earliest stone tool culture.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Orthognathic:<\/strong> The face below the eyes is relatively flat and does not jut out anteriorly.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Paleoanthropologists:<\/strong> Researchers that study human evolution.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Paleoenvironment:<\/strong> An environment from a period in the Earth\u2019s geological past.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Parabolic:<\/strong> Like a parabola (parabola-shaped).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Phalanges:<\/strong> Long bones in the hand and fingers.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Phylogenetics:<\/strong> The study of phylogeny.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Phylogeny:<\/strong> The study of the evolutionary relationships between groups of organisms.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Pliocene:<\/strong> A geological epoch between the Miocene and Pleistocene.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Polytypic:<\/strong> In reference to taxonomy, having two or more group variants capable of interacting and breeding biologically but having morphological population differences.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Postcranium:<\/strong> The skeleton below the cranium (head).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Premolars:<\/strong> The smallest of the hind teeth, behind the canines.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Procumbent:<\/strong> In reference to incisors, tilting forward.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Prognathic:<\/strong> In reference to the face, the area below the eyes juts anteriorly.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Quaternary Ice Age:<\/strong> The most recent geological time period, which includes the Pleistocene and Holocene Epochs and which is defined by the cyclicity of increasing and decreasing ice sheets at the poles.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Relative dating:<\/strong> Dating techniques that refer to a temporal sequence (i.e., older or younger than others in the reference) and do not estimate actual or absolute dates.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Robust:<\/strong> Rugged or exaggerated features.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Site:<\/strong> A place in which evidence of past societies\/species\/activities may be observed through archaeological or paleontological practice.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Specialist:<\/strong> A specialist species can thrive only in a narrow range of environmental conditions or has a limited diet.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Splitters:<\/strong> Researchers who prefer to split a highly variable taxon into multiple groups or species.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Taxa:<\/strong> Plural of taxon, a taxonomic group such as species, genus, or family.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Taxonomy:<\/strong> The science of grouping and classifying organisms.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Techno-complex:<\/strong> A term encompassing multiple assemblages that share similar traits in terms of artifact production and morphology.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Thermoregulation:<\/strong> Maintaining body temperature through physiologically cooling or warming the body.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Ungulates:<\/strong> Hoofed mammals\u2014e.g., cows and kudu.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Volcanic tufts:<\/strong> Rock made from ash from volcanic eruptions in the past.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Valgus knee:<\/strong> The angle of the knee between the femur and tibia, which allows for weight distribution to be angled closer to the point above the center of gravity (i.e., between the feet) in bipeds.<\/span><\/p>\n<h2 class=\"import-Normal\"><strong><span style=\"color: #000000\">About the Authors<br \/>\n<\/span><\/strong><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image31.jpg\" alt=\"A woman with short blonde hair smiles at the camera.\" width=\"311\" height=\"311\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Kerryn Warren, Ph.D.<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Grad Coach International, <a class=\"rId245\" style=\"color: #000000\" href=\"mailto:kerryn.warren@gmail.com\">kerryn.warren@gmail.com<\/a><\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kerryn Warren is a dissertation coach at Grad Coach International and is passionate about stimulating research thinking in students of all levels. She has lectured on multiple topics, including archaeology and human evolution, with her research and science communication interests including hybridization in the hominin fossil record (stemming from research from her Ph.D.) and understanding how evolution is taught in South African schools. She also worked as one of the \u201cUnderground Astronauts,\u201d selected to excavate <em>Homo naledi <\/em>remains from the Rising Star Cave System in the Cradle of Humankind.<\/span><\/p>\n<p>&nbsp;<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image11-2.jpg\" alt=\"A woman with short brown hair smiles at the camera.\" width=\"312\" height=\"306\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">K. Lindsay Hunter, M.A., Ph.D. candidate<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">CARTA, k.lindsay.hunter@gmail.com<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lindsay Hunter is a trained palaeoanthropologist who uses her more than 15 years of experience to make sense of the distant past of our species to build a better future. She received her master\u2019s degree in biological anthropology from the University of Iowa and is completing her Ph.D. in archaeology at the University of the Witwatersrand in Johannesburg, South Africa. She has studied fossil and human bone collections across five continents with major grant support from the National Science Foundation (United States) and the Wenner-Gren Foundation for Anthropological Research. As a National Geographic Explorer, Lindsay developed and managed the National Geographic\u2013sponsored Umsuka Public Palaeoanthropology Project in the Cradle of Humankind World Heritage Site (CoH WHS) in South Africa from within Westbury Township, Johannesburg, between 2016\u20132019. She currently serves as the Community Engagement &amp; Advancement Director for CARTA: The UC San Diego\/Salk Institute Center for Academic Research and Training in Anthropogeny in La Jolla, California.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image25-2.jpg\" alt=\"A woman with black hair stands in a hole in the ground.\" width=\"295\" height=\"339\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Navashni Naidoo, M.Sc.<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">University of Cape Town, nnaidoo2@illinois.edu<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Navashni Naidoo is a researcher at Nelson Mandela University, lecturing on physical geology. She completed her Master\u2019s in Science in Archaeology in 2017 at the University of Cape Town. Her research interests include developing paleoenvironmental proxies suited to the African continent, behavioral ecology, and engaging with community-driven archaeological projects. She has excavated at Stone Age sites across Southern Africa and East Africa. Navashni is currently pursuing a PhD in the Department of Anthropology at the University of Illinois.<\/span><\/p>\n<p>&nbsp;<\/p>\n<p>&nbsp;<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image47.jpg\" alt=\"A man with black hair and dark brown eyes looks at the camera. \" width=\"294\" height=\"294\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Silindokuhle Mavuso, M.Sc.<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">University of Witwatersrand, S.muvaso@ru.ac.za<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Silindokuhle has always been curious about the world around him and how it has been shaped. He is a lecturer at Rhodes University of Witwatersrand (Wits), and conducts research on palaeoenvironmental reconstruction and change of the northeastern Turkana Basin\u2019s Pleistocene sequence. Silindokuhle began his education with a B.Sc. (Geology, Archaeology, and Environmental and Geographical Sciences) from the University of Cape Town before moving to Wits for a B.Sc. Honors (geology and paleontology) and M.Sc. in geology. He is currently concluding his PhD Studies. During this time, he has gained more training as a Koobi Fora Fieldschool fellow (Kenya) as well as an Erasmus Mundus scholar (France). Silindokuhle is a Plio-Pleistocene geologist with a specific focus on identifying and explaining past environments that are associated with early human life and development through time. He is interested in a wide range of disciplines such as micromorphology, sedimentology, geochemistry, geochronology, and sequence stratigraphy. He has worked with teams from significant eastern and southern African hominid sites including Elandsfontein, Rising Star, Sterkfontein, Gondolin, Laetoli, Olduvai, and Koobi Fora.<\/span><\/p>\n<h2 class=\"import-Normal\"><strong><span style=\"color: #000000\">For Further Exploration<br \/>\n<\/span><\/strong><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/humanorigins.si.edu\/evidence\">The Smithsonian Institution website<\/a> hosts descriptions of fossil species, an interactive timeline, and much more.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.maropeng.co.za\/content\/page\/human-evolution\">The Maropeng Museum website<\/a> hosts a wealth of information regarding South African Fossil Bearing sites in the Cradle of Humankind<strong>.<\/strong><\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/perot-museum.imgix.net\/2019-08-naledi-sediba-quick-comparison.pdf\">This quick comparison between <em>Homo naledi<\/em> and <em>Australopithecus sediba<\/em><\/a> from the Perot Museum.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.dropbox.com\/s\/l1d2hv42psj21y9\/Braided%20Stream-1920.mp4?dl=0\">This explanation of the braided stream<\/a> by the Perot Museum.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.hetmp.com\/\">A collation of 3-D files for visualizing<\/a> (or even 3-D printing) for homes, schools, and universities.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.pbslearningmedia.org\/resource\/tdc02.sci.life.evo.lp_humanevo\/human-evolution.\">PBS learning materials<\/a>, including videos and diagrams of the Laetoli footprints, bipedalism, and fossils.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A wealth of <a href=\"https:\/\/australianmuseum.net.au\/learn\/science\/human-evolution\/\">information from the Australian Museum website<\/a>, including species descriptions, family trees, and explanations of bipedalism and diet<strong>.<\/strong><\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\"><strong>References<\/strong><\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Alemseged, Zeresenay, Fred Spoor, William H. Kimbel, Ren\u00e9 Bobe, Denis Geraads, Denn\u00e9 Reed, and Jonathan G. Wynn. 2006. \u201cA Juvenile Early Hominin Skeleton from Dikika, Ethiopia.\u201d <em>Nature<\/em> 443 (7109): 296\u2013301.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Asfaw, Berhane, Tim White, Owen Lovejoy, Bruce Latimer, Scott Simpson, and Gen Suwa. 1999. \u201c<em>Australopithecus garhi<\/em>: A New Species of Early Hominid from Ethiopia.\u201d <em>Science<\/em> 284 (5414): 629\u2013635.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Behrensmeyer, Anna K., Nancy E. Todd, Richard Potts, and Geraldine E. McBrinn. 1997. \u201cLate Pliocene Faunal Turnover in the Turkana Basin, Kenya, and Ethiopia.\u201d <em>Science<\/em> 278 (5343): 637\u2013640.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Berger, Lee R., Darryl J. De Ruiter, Steven E. Churchill, Peter Schmid, Kristian J. Carlson, Paul HGM Dirks, and Job M. Kibii. 2010. \u201c<em>Australopithecus sediba<\/em>: A New Species of <em>Homo<\/em>-like Australopith from South Africa.\u201d <em>Science<\/em> 328 (5975): 195\u2013204.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Bobe, Ren\u00e9, and Anna K. Behrensmeyer. 2004. \u201cThe Expansion of Grassland Ecosystems in Africa in Relation to Mammalian Evolution and the Origin of the Genus <em>Homo<\/em>.\u201d <em>Palaeogeography, Palaeoclimatology, Palaeoecology<\/em> 207 (3\u20134): 399\u2013420.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Brain, C. K. 1967. \u201cThe Transvaal Museum's Fossil Project at Swartkrans.\u201d <em>South African Journal of Science<\/em> 63 (9): 378\u2013384.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1938a. \u201cMore Discoveries of Australopithecus.\u201d <em>Nature<\/em> 141 (1): 828\u2013829.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1938b. \u201cThe Pleistocene Anthropoid Apes of South Africa.\u201d <em>Nature<\/em> 142 (3591): 377\u2013379.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1947. \u201cDiscovery of a New Skull of the South African Ape-Man, Plesianthropus.\u201d <em>Nature<\/em> 159 (4046): 672.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1950. \u201cThe Genera and Species of the South African Fossil Ape-Man.\u201d <em>American Journal of Physical Anthropology<\/em> 8 (1): 1\u201314.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Brunet, Michel, Alain Beauvilain, Yves Coppens, Emile Heintz, Aladji HE Moutaye, and David Pilbeam. 1995. \u201cThe First Australopithecine 2,500 Kilometers West of the Rift Valley (Chad).\u201d <em>Nature<\/em> 378 (6554): 275\u2013273.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Cerling, Thure E., Jonathan G. Wynn, Samuel A. Andanje, Michael I. Bird, David Kimutai Korir, Naomi E. Levin, William Mace, Anthony N. Macharia, Jay Quade, and Christopher H. Remien. 2011. \u201cWoody Cover and Hominin Environments in the Past 6 Million Years.\u201d <em>Nature<\/em> 476, no. 7358 (2011): 51-56..<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clarke, Ronald J. 1998. \u201cFirst Ever Discovery of a Well-Preserved Skull and Associated Skeleton of <em>Australopithecus<\/em>.\u201d <em>South African Journal of Science<\/em> 94 (10): 460\u2013463.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Clarke, Ronald J. 2013. \u201cAustralopithecus from Sterkfontein Caves, South Africa.\u201d In <em>The Paleobiology of Australopithecus<\/em>, edited by K. E. Reed, J. G. Fleagle, and R. E. Leakey, 105\u2013123. Netherlands: Springer.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clarke, Ronald J., and Kathleen Kuman. 2019. \u201cThe Skull of StW 573, a 3.67 Ma Australopithecus Prometheus Skeleton from Sterkfontein Caves, South Africa.\u201d <em>Journal of <\/em><em>H<\/em><em>uman <\/em><em>E<\/em><em>volution<\/em> 134: 102634.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clarke, R. J., and P. V. Tobias. 1995. \u201cSterkfontein Member 2 Foot Bones of the Oldest South African Hominid.\u201d <em>Science<\/em> 269 (5223): 521\u2013524.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Constantino, P. J., and B. A. Wood. 2004. \u201cParanthropus Paleobiology\u201d. In <em>Miscelanea en <\/em><em>Homenae<\/em><em> a Emiliano Aguirre<\/em><em>,<\/em> <em>v<\/em><em>olumen III: Paleoantropologia<\/em>, edited by E. G. P\u00e9rez and S. R. Jara, 136\u2013151. Alcal\u00e1 de Henares: Museo Arqueologico Regional.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Constantino, P. J., and B. A. Wood. 2007. \u201cThe Evolution of Zinjanthropus boisei.\u201d <em>Evolutionary Anthropology: <\/em><em>Issues, News, and Reviews<\/em> 16 (2): 49\u201362.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Dart, Raymond A. 1925. \u201cAustralopithecus africanus, the Man-Ape of South Africa.\u201d <em>Nature<\/em> 115: 195\u2013199.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Darwin, Charles. 1871. <em>The Descent of Man: And Selection in Relation to Sex<\/em>. London: J. Murray.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Daver, Guillaume, F. Guy, Hassane Ta\u00efsso Mackaye, Andossa Likius, J-R. Boisserie, Abderamane Moussa, Laurent Pallas, Patrick Vignaud, and N\u00e9koulnang D. Clarisse. 2022. \"Postcranial Evidence of Late Miocene Hominin Bipedalism in Chad.\" <em>Nature<\/em> 609 (7925): 94\u2013100.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Heinzelin, Jean de, J. Desmond Clark, Tim White, William Hart, Paul Renne, Giday WoldeGabriel, Yonas Beyene, and Elisabeth Vrba. 1999. \u201cEnvironment and Behavior of 2.5-Million-Year-Old Bouri Hominids.\u201d <em>Science<\/em> 284 (5414): 625\u2013629.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">DeMenocal, Peter B. D. 2004. \u201cAfrican Climate Change and Faunal Evolution during the Pliocene\u2013Pleistocene.\u201d <em>Earth and Planetary Science Letters<\/em> 220 (1\u20132): 3\u201324.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">DeMenocal, Peter B. D. and J. Bloemendal, J. 1995. \u201cPlio-Pleistocene Climatic Variability in Subtropical Africa and the Paleoenvironment of Hominid Evolution: A Combined Data-Model Approach.\u201d In <em>Paleoclimate and Evolution, with Emphasis on Human Origins<\/em>, edited by E. S. Vrba, G. H. Denton, T. C. Partridge, and L. H. Burckle, 262\u2013288. New Haven: Yale University Press.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Dirks, Paul HGM, Job M. Kibii, Brian F. Kuhn, Christine Steininger, Steven E. Churchill, Jan D. Kramers, Robyn Pickering, Daniel L. Farber, Anne-Sophie M\u00e9riaux, Andy I. R. Herries, Geoffrey C. P. King, And Lee R. Berger. 2010. \u201cGeological Setting and Age of <em>Australopithecus sediba<\/em> from Southern Africa.\u201d <em>Science<\/em> 328 (5975): 205\u2013208.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Faith, J. Tyler, and Anna K. Behrensmeyer. 2013. \u201cClimate Change and Faunal Turnover: Testing the Mechanics of the Turnover-Pulse Hypothesis with South African Fossil Data.\u201d <em>Paleobiology<\/em> 39 (4): 609\u2013627.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Grine, Frederick E. 1988. \u201cNew Craniodental Fossils of <em>Paranthropus<\/em> from the Swartkrans Formation and Their Significance in \u2018Robust\u2019 Australopithecine Evolution.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by F. E. Grine, 223\u2013243. New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Grine, Frederick E., Carrie S. Mongle, John G. Fleagle, and Ashley S. Hammond. 2022. \"The Taxonomic Attribution of African Hominin Postcrania from the Miocene through the Pleistocene: Associations and Assumptions.\" <em>Journal of Human Evolution<\/em> 173: 103255.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Haile-Selassie, Yohannes, Luis Gibert, Stephanie M. Melillo, Timothy M. Ryan, Mulugeta Alene, Alan Deino, Naomi E. Levin, Gary Scott, and Beverly Z. Saylor. 2015. \u201cNew Species from Ethiopia Further Expands Middle Pliocene Hominin Diversity.\u201d <em>Nature<\/em> 521 (7553): 432\u2013433.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Haile-Selassie, Yohannes, Stephanie M. Melillo, Antonino Vazzana, Stefano Benazzi, and Timothy M. Ryan. 2019. \u201cA 3.8-Million-Year-Old Hominin Cranium from Woranso-Mille, Ethiopia.\u201d <em>Nature<\/em> 573 (7773): 214-219.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Harmand, Sonia, Jason E. Lewis, Craig S. Feibel, Christopher J. Lepre, Sandrine Prat, Arnaud Lenoble, Xavier Bo\u00ebs et al. 2015. \u201c3.3-Million-Year-Old Stone Tools from Lomekwi3, West Turkana, Kenya.\u201d <em>Nature<\/em> 521 (7552): 310\u2013316.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Hay, Richard L. 1990. \u201cOlduvai Gorge: A Case History in the Interpretation of Hominid Paleoenvironments.\u201d In <em>East Africa: Establishment of a Geologic Framework for Paleoanthropology<\/em>, edited by L. Laporte, 23\u201337<em>.<\/em> Boulder: Geological Society of America.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Hay, Richard L., and Mary D. Leakey. 1982. \u201cThe Fossil Footprints of Laetoli.\u201d <em>Scientific American<\/em> 246 (2): 50\u201357.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Hlazo, Nomawethu. 2015. \u201cParanthropus: Variation in Cranial Morphology.\u201d Honours thesis, Archaeology Department, University of Cape Town, Cape Town.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Hlazo, Nomawethu. 2018. \u201cVariation and the Evolutionary Drivers of Diversity in the Genus <em>Paranthropus<\/em>.\u201d Master\u2019s thesis, Archaeology Department, University of Cape Town, Cape Town.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Johanson, D. C., T. D. White, and Y. Coppens. 1978. \u201cA New Species of the Genus <em>Australopithecus<\/em> (Primates: Hominidae) from the Pliocene of East Africa.\u201d <em>Kirtlandia<\/em> 28: 1\u201314.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kimbel, William H. 2015. \u201cThe Species and Diversity of Australopiths.\u201d In <em>Handbook of Paleoanthropology<\/em>, 2nd ed., edited by T. Hardt, 2071\u20132105. Berlin: Springer.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kimbel, William H., and Lucas K. Delezene. 2009. \u201c\u2018Lucy\u2019 Redux: A Review of Research on <em>Australopithecus afarensis<\/em>.\u201d <em>American <\/em><em>J<\/em><em>ournal of <\/em><em>P<\/em><em>hysical <\/em><em>A<\/em><em>nthropology<\/em> 140 (S49): 2\u201348.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Kingston, John D. 2007. \u201cShifting Adaptive Landscapes: Progress and Challenges in Reconstructing Early Hominid Environments.\u201d <em>American Journal of Physical Anthropology<\/em> 134 (S45): 20\u201358.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Kingston, John D., and Terry Harrison. 2007. \u201cIsotopic Dietary Reconstructions of Pliocene Herbivores at Laetoli: Implications for Early Hominin Paleoecology.\u201d <em>Palaeogeography, Palaeoclimatology, Palaeoecology<\/em> 243 (3\u20134): 272\u2013306.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Leakey, Louis S. B. 1959. \u201cA New Fossil Skull from Olduvai.\u201d <em>Nature<\/em> 184 (4685): 491\u2013493.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Leakey, Mary 1971. <em>Olduvai Gorge<\/em>, Vol. 3. Cambridge: Cambridge University Press.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Leakey, Mary D., and Richard L. Hay. 1979. \u201cPliocene Footprints in the Laetoli Beds at Laetoli, Northern Tanzania.\u201d <em>Nature<\/em> 278 (5702): 317\u2013323.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Leakey, Meave G., Craig S. Feibel, Ian McDougall, and Alan Walker. 1995. \u201cNew Four\u2013Million-Year-Old Hominid Species from Kanapoi and Allia Bay, Kenya.\u201d <em>Nature<\/em> 376 (6541): 565\u2013571.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Meave G., Fred Spoor, Frank H. Brown, Patrick N. Gathogo, Christopher Kiarie, Louise N. Leakey, and Ian McDougall. 2001. \u201cNew Hominin Genus from Eastern Africa Shows Diverse Middle Pliocene Lineages.\u201d <em>Nature<\/em> 410 (6827): 433\u2013440.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lebatard, Anne-Elisabeth, Didier L. Bourl\u00e8s, Philippe Duringer, Marc Jolivet, R\u00e9gis Braucher, Julien Carcaillet, Mathieu Schuster et al. 2008. \u201cCosmogenic Nuclide Dating of <em>Sahelanthropus tchadensis<\/em> and <em>Australopithecus bahrelghazali<\/em>: Mio-Pliocene Hominids from Chad.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 105 (9): 3226\u20133231.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lee-Thorp, Julia. 2011. \u201cThe Demise of \u2018Nutcracker Man.\u2019\u201d <em>Proceedings of the National Academy of Sciences<\/em> 108 (23): 9319\u20139320.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lombard, Marlize, L. Y. N. Wadley, Janette Deacon, Sarah Wurz, Isabelle Parsons, Moleboheng Mohapi, Joane Swart, and Peter Mitchell. 2012. \u201cSouth African and Lesotho Stone Age Sequence Updated.\u201d <em>The South African Archaeological Bulletin<\/em> 67 (195): 123\u2013144.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Maslin, Mark A., Chris M. Brierley, Alice M. Milner, Susanne Shultz, Martin H. Trauth, and Katy E. Wilson. 2014. \u201cEast African Climate Pulses and Early Human Evolution.\u201d <em>Quaternary Science Reviews<\/em> 101: 1\u201317.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">McHenry, Henry M. 2009. \u201cHuman Evolution.\u201d In <em>Evolution: The First Four Billion Years<\/em>, edited by M. Ruse and J. Travis, 256\u2013280. Cambridge: The Belknap Press of Harvard University Press..<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Patterson, Bryan, and William W. Howells. 1967. \u201cHominid Humeral Fragment from Early Pleistocene of Northwestern Kenya.\u201d <em>Science<\/em> 156 (3771): 64\u201366.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Pickering, Robyn, and Jan D. Kramers. 2010. \u201cRe-appraisal of the Stratigraphy and Determination of New U-Pb Dates for the Sterkfontein Hominin Site.\u201d <em>Journal of Human Evoluti<\/em><em>on<\/em> 59 (1): 70\u201386.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Potts, Richard. 1998. \u201cEnvironmental Hypotheses of Hominin Evolution.\u201d <em>American Journal of Physical Anthropology<\/em> 107 (S27): 93\u2013136.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Potts, Richard. 2013. \u201cHominin Evolution in Settings of Strong Environmental Variability.\u201d <em>Quaternary Science Reviews<\/em> 73: 1\u201313.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Rak, Yoel. 1983. <em>The <\/em><em>A<\/em><em>ustralopithecine <\/em><em>F<\/em><em>ace<\/em>. New York: Academic Press.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Rak, Yoel. 1988. \u201cOn Variation in the Masticatory System of <em>Australopithecus boisei<\/em>.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by M. Ruse and J. Travis, 193\u2013198<em>.<\/em> New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Semaw, Sileshi. 2000. \u201cThe World\u2019s Oldest Stone Artefacts from Gona, Ethiopia: Their Implications for Understanding Stone Technology and Patterns of Human Evolution between 2.6 Million Years Ago and 1.5 Million Years Ago.\u201d <em>Journal of Archaeological Science<\/em> 27(12): 1197\u20131214.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Shipman, Pat. 2002. <em>The Man Who Found the Missing Link: Eug<\/em><em>e<\/em><em>ne Dubois and <\/em><em>h<\/em><em>is Lifelong Quest to Prove Darwin Right<\/em>. New York: Simon &amp; Schuster.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Spoor, Fred. 2015. \u201cPalaeoanthropology: The Middle Pliocene Gets Crowded.\u201d<em> Nature<\/em> 521 (7553): 432\u2013433.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Strait, David S., Frederick E. Grine, and Marc A. Moniz. 1997. A Reappraisal of Early Hominid Phylogeny.\u201d <em>Journal of <\/em><em>H<\/em><em>uman <\/em><em>E<\/em><em>volution<\/em> 32 (1): 17\u201382.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Thackeray, J. Francis. 2000. \u201c\u2018Mrs. Ples\u2019 from Sterkfontein: Small Male or Large Female?\u201d <em>The South African Archaeological <\/em><em>Bulletin<\/em> 55: 155\u2013158.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Thackeray, J. Francis, Jos\u00e9 Braga, Jacques Treil, N. Niksch, and J. H. Labuschagne. 2002. \u201c\u2018Mrs. Ples\u2019 (Sts 5) from Sterkfontein: An Adolescent Male?\u201d <em>South African Journal of Science<\/em> 98 (1\u20132): 21\u201322.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Toth, Nicholas. 1985. \u201cThe Oldowan Reassessed.\u201d <em>Journal of Archaeological Science<\/em>\u00a012 (2): 101\u2013120.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Vrba, E. S. 1988. \u201cLate Pliocene Climatic Events and Hominid Evolution.\u201d In <em>The <\/em><em>E<\/em><em>volutionary <\/em><em>H<\/em><em>istory of the <\/em><em>R<\/em><em>obust Australopithecines<\/em>, edited by F. E. Grine, 405\u2013426. New York: Aldine.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Vrba, Elisabeth S. 1998. \u201cMultiphasic Growth Models and the Evolution of Prolonged Growth Exemplified by Human Brain Evolution.\u201d <em>Journal of Theoretical Biology<\/em> 190 (3): 227\u2013239.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Vrba, Elisabeth S. 2000. \u201cMajor Features of Neogene Mammalian Evolution in Africa.\u201d In <em>Cenozoic <\/em><em>G<\/em><em>eology of <\/em><em>S<\/em><em>outhern Africa<\/em>, edited by T. C. Partridge and R. Maud, 277\u2013304<em>.<\/em> Oxford: Oxford University Press.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Walker, Alan C., and Richard E. Leakey. 1988. \u201cThe Evolution of <em>Australopithecus boisei<\/em>.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by F. E. Grine, 247\u2013258. New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Walker, Alan, Richard E. Leakey, John M. Harris, and Francis H. Brown. 1986. \u201c2.5-my <em>Australopithecus boisei<\/em> from West of Lake Turkana, Kenya.\u201d <em>Nature<\/em> 322 (6079): 517\u2013522.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Ward, Carol, Meave Leakey, and Alan Walker. 1999. \u201cThe New Hominid Species <em>Australopithecus anamensis<\/em>.\u201d <em>Evolutionary Anthropology<\/em> 7 (6): 197\u2013205.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">White, Tim D. 1988. \u201cThe Comparative Biology of \u2018Robust\u2019 Australopithecus: Clues from Content.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by F. E. Grine, 449\u2013483. New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">White, Tim D., Gen Suwa, and Berhane Asfaw. 1994. \u201c<em>Australopithecus ramidus<\/em>, a New Species of Early Hominid from Aramis, Ethiopia.\u201d <em>Nature<\/em> 371 (6495): 306\u2013312.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Wood, Bernard. 2010. \u201cReconstructing Human Evolution: Achievements, Challenges, and Opportunities.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 10 (2): 8902\u20138909.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Wood, Bernard, and Eve K. Boyle. 2016. \u201cHominin Taxic Diversity: Fact or Fantasy?\u201d <em>Yearbook of Physical Anthropology<\/em> 159 (S61): 37\u201378.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Wood, Bernard, and Kes Schroer. 2017. \u201cParanthropus: Where Do Things Stand?\u201d In <em>Human Paleontology and Prehistory<\/em>, edited by A. Marom and E. Hovers, 95\u2013107. New York: Springer, Cham.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Acknowledgements<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">All of the authors in this section are students and early career researchers in paleoanthropology and related fields in South Africa (or at least have worked in South Africa). We wish to thank everyone who supports young and diverse talent in this field and would love to further acknowledge Black, African, and female academics who have helped pave the way for us.<\/span><\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1766\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1766\"><div tabindex=\"-1\"><p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kerryn Warren, Ph.D., Grad Coach International<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lindsay Hunter, M.A., University of Iowa<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Navashni Naidoo, M.Sc., University of Cape Town<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Silindokuhle Mavuso, M.Sc., University of Witwatersrand<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\"><em>This chapter is a revision from <\/em><em>\"<\/em><a class=\"rId7\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\"><em>Chapter 9: Early Hominins<\/em><\/a><em>\" <\/em><em>by Kerryn Warren, K. Lindsay Hunter, Navashni Naidoo, Silindokuhle Mavuso, Kimberleigh Tommy, Rosa Moll, and Nomawethu Hlazo<\/em><em>. In <\/em><a class=\"rId8\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, first edition<\/em><\/a><em>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under <\/em><a class=\"rId9\" style=\"color: #000000\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\"><em>CC BY-NC 4.0<\/em><\/a><em>. <\/em><\/span><\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\"><span style=\"color: #000000\"><span style=\"color: #ffffff\">Learning Objectives<\/span><br \/>\n<\/span><\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li><span style=\"color: #000000\">Understand what is meant by \u201cderived\u201d and \u201cancestral\u201d traits and why this is relevant for understanding early hominin evolution.<\/span><\/li>\n<li><span style=\"color: #000000\">Understand changing paleoclimates and paleoenvironments as potential factors influencing early hominin adaptations.<\/span><\/li>\n<li><span style=\"color: #000000\">Describe the anatomical changes associated with bipedalism and dentition in early hominins, as well as their implications..<\/span><\/li>\n<li><span style=\"color: #000000\">Describe early hominin genera and species, including their currently understood dates and geographic expanses.<\/span><\/li>\n<li><span style=\"color: #000000\">Describe the earliest stone tool techno-complexes and their impact on the transition from early hominins to our genus.<\/span><\/li>\n<\/ul>\n<\/div>\n<\/div>\n<h2 class=\"__UNKNOWN__\"><span style=\"color: #000000\">Defining Hominins<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">It is through our study of our hominin ancestors and relatives that we are exposed to a world of \u201cmight have beens\u201d: of other paths not taken by our species, other ways of being human. But to better understand these different evolutionary trajectories, we must first define the terms we are using. If an imaginary line were drawn between ourselves and our closest relatives, the great apes, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1200\">bipedalism<\/a><\/strong> (or habitually walking upright on two feet) is where that line would be. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1270\">Hominin<\/a><\/strong>, then, means everyone on \u201cour\u201d side of the line: humans and all of our extinct bipedal ancestors and relatives since our divergence from the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1272\">last common ancestor (LCA)<\/a><\/strong> we share with chimpanzees.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Historic interpretations of our evolution, prior to our finding of early hominin <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1274\"><strong>fossils<\/strong><\/a>, varied. Debates in the mid-1800s regarding hominin origins focused on two key issues:<\/span><\/p>\n<ul>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">Where did we evolve?<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">Which traits evolved first?<\/span><\/li>\n<\/ul>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff00ff\">Charles Darwin hypothesized that we evolved in Africa, as he was convinced that we shared greater commonality with chimpanzees and gorillas on the continent (Darwin 1871). Others, such as Ernst Haeckel and Eug\u00e8ne Dubois, insisted that we were closer in affinity to orangutans and that we evolved in Eurasia where, until the discovery of the Taung Child in South Africa in 1924, all humanlike fossils (of Neanderthals and <em>Homo erectus<\/em>) had been found (Shipman 2002).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Within this conversation, naturalists and early <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1276\">paleoanthropologists<\/a><\/strong> (people who study human evolution) speculated about which human traits came first. These included the evolution of a big brain (<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1278\"><strong>encephalization<\/strong><\/a>), the evolution of the way in which we move about on two legs (bipedalism), and the evolution of our flat faces and small teeth (indications of dietary change). Original hypotheses suggested that, in order to be motivated to change diet and move about in a bipedal fashion, the large brain needed to have evolved first, as is seen in the fossil species mentioned above.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">However, we now know that bipedal locomotion is one of the first things that evolved in our lineage, with early relatives having more apelike dentition and small brain sizes. While brain size expansion is seen primarily in our genus, <em>Homo<\/em>, earlier hominin brain sizes were highly variable between and within taxa, from 300 cc (cranial capacity, cm<sup>3<\/sup>), estimated in <em>Ardipithecus<\/em>, to 550 cc, estimated in <em>Paranthropus boisei<\/em>. The lower estimates are well within the range of variation of nonhuman extant great apes. In addition, body size variability also plays a role in the interpretation of whether brain size could be considered large or small for a particular species or specimen. In this chapter, we will tease out the details of early hominin evolution in terms of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1280\"><strong>morphology<\/strong><\/a> (i.e. the study of the form, size, or shape of things; in this case, skeletal parts).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">We also know that early human evolution occurred in a very complicated fashion. There were multiple species (multiple genera) that featured diversity in their diets and locomotion. Specimens have been found all along the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1282\"><strong>East African Rift System <\/strong>(<strong>EARS)<\/strong><\/a>; that is, in Ethiopia, Kenya, Tanzania, and Malawi; see Figure 9.1), in limestone caves in South Africa, and in Chad. Dates of these early relatives range from around 7 million years ago (mya) to around 1 mya, overlapping temporally with members of our genus, <em>Homo<\/em>.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 610px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/06\/image38.png\" alt=\"Patchy green mountain alongside a deep sandy valley in East Africa.\" width=\"610\" height=\"277\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.1: East African Rift System (EARS). Credit: <a href=\"https:\/\/www.flickr.com\/photos\/ninara\/8624605781\/in\/photolist-x2yH7-x2yHe-VfVWuD-e98mPF-SzzjsU-2bsBZhC-2hHec7m-xtJ7Ez-NXnXvh-7Yg3uo-2cS3FgG-2hjo1Dc-2hjGoTS-nnumi8-82U66W-dMNn7B-8jdVbd-NWDg8-NW6fj-ebhx5w-bkFv1G-Ct5ZD-5JQk8A-y6TgAc-x9k6oe-2ebLTDC-WcPMnJ-2ekh6CS-Cu3LH-xNHDFK-9RUsZi-94jVt4-P46uiB-QFyjyE-crU8N7-5JLJKV-2ekSgk8-5JL454-2cPgZrF-2bHfQZu-dMTVPN-6yUbeN-jzMicQ-48XjU9-2etR2Ze-Styrvw-crU7V7-2wakq3-crU6Z1-2etR2XR\/\">IMG_1696 Great Rift Valley<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/ninara\/\">Ninara<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\">CC BY 2.0 License<\/a>.<\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Yet there is still so much to understand. Modern debates now look at the relatedness of these species to us and to one another, and they consider which of these species were able to make and use tools. As a result, every <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1284\">site<\/a><\/strong> discovery in the patchy hominin fossil record tells us more about our evolution. In addition, recent scientific techniques (not available even ten years ago) provide new insights into the diets, environments, and lifestyles of these ancient relatives.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In the past, <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1288\"><strong>taxonom<\/strong><strong>y<\/strong><\/a> was primarily based on morphology. Today it is tied to known relationships based on molecular <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1290\"><strong>phylogeny<\/strong><\/a> (e.g., based on DNA) or a combination of the two. This is complicated when applied to living <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1286\"><strong>taxa<\/strong><\/a>, but becomes much more difficult when we try to categorize ancestor-descendant relationships for long-extinct species whose molecular information is no longer preserved. We therefore find ourselves falling back on morphological comparisons, often of teeth and partially fossilized skeletal material.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">It is here that we turn to the related concepts of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1292\">cladistics<\/a> <\/strong>and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1294\">phylogenetics<\/a><\/strong><strong>. <\/strong><span style=\"background-color: #ff99cc\">Cladistics groups organisms according to their last common ancestors based on shared <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1126\">derived traits<\/a><\/strong>. <\/span>In the case of early hominins, these are often morphological traits that differ from those seen in earlier populations. These new or modified traits provide evidence of evolutionary relationships, and organisms with the same derived traits are grouped in the same <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1116\">clade<\/a> <\/strong>(Figure 9.2). <span style=\"background-color: #ff99cc\">For example, if we use feathers as a trait, we can group pigeons and ostriches into the clade of birds.<\/span> In this chapter, we will examine the grouping of the Robust Australopithecines, whose cranial and dental features differ from those of earlier hominins, and therefore are considered derived.<\/span><\/p>\n<figure style=\"width: 708px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image48.png\" alt=\"Phylogenetic tree shows clades and non clade groupings.\" width=\"708\" height=\"192\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.2: Clades refer to groups of species or taxa that share a common ancestor. In <span class=\"ILfuVd\" lang=\"en\"><span class=\"hgKElc\">a phylogeny, a clade is a complete group of lineages, including their last common ancestor. Groupings that do not include a common ancestor and <em>all<\/em> of its descendants are not clades. <\/span><\/span>Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\">Clades (Figure 9.2)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Katie Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Dig Deeper: Problems Defining Hominin Species<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000\">It is worth noting that species designations for early hominin specimens are often highly contested. This is due to the fragmentary nature of the fossil record, the large timescale (millions of years) with which paleoanthropologists need to work, and the difficulty in evaluating whether morphological differences and similarities are due to meaningful phylogenetic or biological differences or subtle differences\/variation in niche occupation or time. In other words, do morphological differences really indicate different species? How would classifying species in the paleoanthropological record compare with classifying living species today, for whom we can sequence genomes and observe lifestyles?<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">There are also broader philosophical differences among researchers when it comes to paleo-species designations. Some scientists, known as \u201c<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1296\"><strong>lumpers<\/strong>,<\/a>\u201d argue that large variability is expected among multiple populations in a given species over time. These researchers will therefore prefer to \u201clump\u201d specimens of subtle differences into single taxa. Others, known as \u201c<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1298\"><strong>splitters<\/strong><\/a>,\u201d argue that species variability can be measured and that even subtle differences can imply differences in niche occupation that are extreme enough to mirror modern species differences. In general, splitters would consider geographic differences among populations as meaning that a species is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1300\">polytypic<\/a><\/strong> <span style=\"background-color: #ff99cc\">(i.e., capable of interacting and breeding biologically but having morphological population differences)<\/span>. This is worth keeping in mind when learning about why species designations may be contested.<\/span><\/p>\n<figure style=\"width: 520px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image8.jpg\" alt=\"A graph shows a curved line depicting changes in morphology among two species over time.\" width=\"520\" height=\"292\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.3: This graph demonstrates the concept of a chronospecies, where one species (Species A) \u201cevolves\u201d into another (Species B). Credit: Chronospecies original to Explorations: An Open Invitation to Biological Anthropology, 2nd edition by Kerryn Warren is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\"><span style=\"color: #000000\">This further plays a role in evaluating ancestry. Debates over which species \u201cgave rise\u201d to which continue to this day. It is common to try to create \u201clineages\u201d of species to determine when one species evolved into another over time. We refer to these as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1302\"><strong>chronospecies<\/strong><\/a> (Figure 9.3). Constructed hominin phylogenetic trees are routinely variable, changing with new specimen discoveries, new techniques for evaluating and comparing species, and, some have argued, nationalist or biased interpretations of the record. More recently, some researchers have shifted away from \u201ctreelike\u201d models of ancestry toward more nuanced metaphors such as the \u201cbraided stream,\u201d where some levels of interbreeding among species and populations are seen as natural processes of evolution.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Finally, it is worth considering the process of fossil discovery and publication. Some fossils are easily diagnostic to a species level and allow for easy and accurate interpretation. Some, however, are more controversial. This could be because they do not easily preserve or are incomplete, making it difficult to compare and place within a specific species (e.g., a fossil of a patella or knee bone). Researchers often need to make several important claims when announcing or publishing a find: a secure date (if possible), clear association with other finds, and an adequate comparison among multiple species (both extant and fossil). Therefore, it is not uncommon that an important find was made years before it is scientifically published.<\/span><\/p>\n<\/div>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Paleoenvironment and Hominin Evolution<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There is no doubt that one of the major selective pressures in hominin evolution is the environment. <span style=\"background-color: #ff9900\"><span style=\"background-color: #ffff00\">Large-scale changes in global and regional climate, as well as alterations to the environment, are<\/span><span style=\"background-color: #ffff00\"> (<\/span><\/span><em style=\"background-color: transparent\"><span style=\"text-decoration: underline\">thought to be<\/span>)<\/em><span style=\"background-color: #ffff00\"><del> all<\/del> linked to (all) hominin diversification, dispersal, and extinction<\/span> (Maslin et al. 2014). Environmental reconstructions often use modern analogues. Let us take, for instance, the hippopotamus. It is an animal that thrives in environments that have abundant water to keep its skin cool and moist. If the environment for some reason becomes drier, it is expected that hippopotamus populations will reduce. If a drier environment becomes wetter, it is possible that hippopotamus populations may be attracted to the new environment and thrive. Such instances have occurred multiple times in the past, and the bones of some <strong style=\"background-color: transparent\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1308\">fauna<\/a><\/strong> (i.e., animals, like the hippopotamus) that are sensitive to these changes give us insights into these events.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Yet reconstructing a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1304\">paleoenvironment<\/a><\/strong> relies on a range of techniques, which vary depending on whether research interests focus on local changes or more global environmental changes\/reconstructions. For local environments (such as a single site or region), comparing the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1306\">faunal assemblages<\/a> <\/strong>(collections of fossils of animals found at a site) with animals found in certain modern environments allows us to determine if past environments mirror current ones in the region. Changes in the faunal assemblages, as well as when they occur and how they occur, tell us about past environmental changes. Other techniques are also useful in this regard. Chemical analyses, for instance, can reveal the diets of individual fauna, providing clues as to the relative wetness or dryness of their environment (e.g., nitrogen <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1310\"><strong>isotopes<\/strong><\/a>; Kingston and Harrison 2007).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Global climatic changes in the distant past, which fluctuated between being colder and drier and warmer and wetter on average, would have global implications for environmental change (Figure 9.4). These can be studied by comparing marine core and terrestrial soil data across multiple sites. These techniques are based on chemical analysis, such as examination of the nitrogen and oxygen isotopes in shells and sediments. Similarly, analyzing pollen grains shows which kinds of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1312\"><strong>flora<\/strong> <\/a> survived in an environment at a specific time period. There are multiple lines of evidence that allow us to visualize global climate trends over millions of years (although it should be noted that the direction and extent of these changes could differ by geographic region).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 649px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image12-1-1.png\" alt=\"Chart shows cyclical carbon dioxide levels from 800,000 years ago until today.\" width=\"649\" height=\"406\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.4: This graph, based on the comparison of atmospheric samples contained in ice cores and more recent direct measurements, illustrates how atmospheric CO\u2082 has fluctuated over time and increased sharply since the Industrial Revolution. The graph also shows that since 800,000ya (and before) atmospheric CO\u2082 has never exceeded 300 parts per million (ppm). In 1950 it was 310ppm. Today atmospheric CO\u2082 has spiked to over 410 ppm. Credit: <a href=\"https:\/\/climate.nasa.gov\/evidence\/\">CO\u2082 increase since the Industrial Revolution<\/a> by <a href=\"https:\/\/www.nasa.gov\/\">NASA<\/a> is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a> and is used within <a href=\"https:\/\/www.nasa.gov\/multimedia\/guidelines\/index.html\">NASA guidelines on re-use<\/a>. Original from Luthi, D., et al.. 2008; Etheridge, D.M., et al. 2010; Vostok ice core data\/J.R. Petit et al.; NOAA Mauna Loa CO<a href=\"https:\/\/climate.nasa.gov\/evidence\/\">\u2082<\/a> record..<\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Both local and global climatic\/environmental changes have been used to understand factors affecting our evolution (DeHeinzelin et al. 1999; Kingston 2007). Environmental change acts as an important factor regarding the onset of several important hominin traits seen in early hominins and discussed in this chapter. Namely, the environment has been interpreted as the following:<\/span><\/p>\n<ul>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">the driving force behind the evolution of bipedalism,<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">the reason for change and variation in early hominin diets, and<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">the diversification of multiple early hominin species.<\/span><\/li>\n<\/ul>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are numerous hypotheses regarding how climate has driven and continues to drive human evolution. Here, we will focus on just three popular hypotheses.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Savannah Hypothesis (or Aridity Hypothesis)<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The hypothesis:<\/strong> This popular theory suggests that the expansion of the savannah (or less densely forested, drier environments) forced early hominins from an <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1314\"><strong>arboreal<\/strong> <\/a> lifestyle (one living in trees) to a terrestrial one where bipedalism was a more efficient form of locomotion (Figure 9.5). It was first proposed by Darwin (1871) and supported by anthropologists like Raymond Dart (1925). However, this idea was supported by little fossil or paleoenvironmental evidence and was later refined as the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1318\">Aridity Hypothesis<\/a><\/strong>. This hypothesis states that the long-term <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1316\">aridification<\/a><\/strong> and, thereby, expansion of savannah biomes were drivers in diversification in early hominin evolution (deMenocal 2004; deMenocal and Bloemendal 1995). It advocates for periods of accelerated aridification leading to early hominin speciation events.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 647px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image46.png\" alt=\"Photograph showing a dry, open savannah environment.\" width=\"647\" height=\"486\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.5: The African savannah grew during early hominin evolution. This may have forced early hominins from an arboreal lifestyle to a terrestrial one, where bipedalism was a more efficient form of locomotion. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:African_savannah_@_Masai_Mara_(21308330314).jpg\">African savannah @ Masai Mara (21308330314)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/132394214@N04\">Leo Li<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\">CC BY 2.0 License<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The evidence:<\/strong> While early bipedal hominins are often associated with wetter, more closed environments (i.e., not the Savannah Hypothesis), both marine and terrestrial records seem to support general cooling, drying conditions, with isotopic records indicating an increase in grasslands (i.e., colder and wetter climatic conditions) between 8 mya and 6 mya across the African continent (Cerling et al. 2011). This can be contrasted with later climatic changes derived from aeolian dust records (sediments transported to the site of interest by wind), which demonstrate increases in seasonal rainfall between 3 mya and 2.6 mya, 1.8 mya and 1.6 mya, and 1.2 mya and 0.8 mya (deMenocal 2004; deMenocal and Bloemendal 1995).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Interpretation(s):<\/strong> Despite a relatively scarce early hominin record, it is clear that two important factors occur around the time period in which we see increasing aridity. The first factor is the diversification of taxa, where high morphological variation between specimens has led to the naming of multiple hominin genera and species. The second factor is the observation that the earliest hominin fossils appear to have traits associated with bipedalism and are dated to around the drying period (as based on isotopic records). Some have argued that it is more accurately a combination of bipedalism and arboreal locomotion, which will be discussed later. However, the local environments in which these early specimens are found (as based on the faunal assemblages) do not appear to have been dry.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Turnover Pulse Hypothesis<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The hypothesis:<\/strong> In 1985, paleontologist Elisabeth Vbra noticed that in periods of extreme and rapid climate change, <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1326\"><strong>ungulates<\/strong><\/a> (hoofed mammals of various kinds) that had generalized diets fared better than those with specialized diets (Vrba 1988, 1998). <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1322\"><strong>Specialist<\/strong><\/a> eaters <span style=\"background-color: #ff99cc\">(those who rely primarily on specific food types)<\/span> faced extinction at greater rates than their <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1320\">generalist<\/a> <\/strong><span style=\"background-color: #ff99cc\">(those who can eat more varied and variable diets) <\/span>counterparts because they were unable to adapt to new environments (Vrba 2000). Thus, periods with extreme climate change would be associated with high <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1324\"><strong>faunal turnover<\/strong><\/a>: that is, the extinction of many species and the speciation, diversification, and migration of many others to occupy various niches.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The evidence:<\/strong> The onset of the<strong> <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1400\">Quaternary Ice Age<\/a><\/strong>, between 2.5 mya and 3 mya, brought extreme global, cyclical <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1328\"><strong>interglacial<\/strong> <\/a> and <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1330\"><strong>glacial<\/strong><\/a> periods (warmer, wetter periods with less ice at the poles, and colder, drier periods with more ice near the poles). Faunal evidence from the Turkana basin in East Africa indicates multiple instances of faunal turnover and extinction events, in which global climatic change resulted in changes from closed\/forested to open\/grassier habitats at single sites (Behrensmeyer et al. 1997; Bobe and Behrensmeyer 2004). Similarly, work in the Cape Floristic Belt of South Africa shows that extreme changes in climate play a role in extinction and migration in ungulates. While this theory was originally developed for ungulates, its proponents have argued that it can be applied to hominins as well. However, the link between climate and speciation is only vaguely understood (Faith and Behrensmeyer 2013).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Interpretation(s):<\/strong> While the evidence of rapid faunal turnover among ungulates during this time period appears clear, there is still some debate around its usefulness as applied to the paleoanthropological record. Specialist hominin species do appear to exist for long periods of time during this time period, yet it is also true that <em>Homo<\/em>, a generalist genus with a varied and adaptable diet, ultimately survives the majority of these fluctuations, and the specialists appear to go extinct.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Variability Selection Hypothesis<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The hypothesis: <\/strong>This hypothesis was first articulated by paleoanthropologist Richard Potts (1998). It links the high amount of climatic variability over the last 7 million years to both behavioral and morphological changes. Unlike previous notions, this hypothesis states that hominin evolution does not respond to habitat-specific changes or to specific aridity or moisture trends. Instead, long-term environmental unpredictability over time and space influenced morphological and behavioral adaptations that would help hominins survive, regardless of environmental context (Potts 1998, 2013). The Variability Selection Hypothesis states that hominin groups would experience varying degrees of natural selection due to continually changing environments and potential group isolation. This would allow certain groups to develop genetic combinations that would increase their ability to survive in shifting environments. These populations would then have a genetic advantage over others that were forced into habitat-specific adaptations (Potts 2013).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The evidence:<\/strong> The evidence for this theory is similar to that for the Turnover Pulse Hypothesis: large climatic variability and higher survivability of generalists versus specialists. However, this hypothesis accommodates for larger time-scales of extinction and survival events.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Interpretation(s):<\/strong> In this way, the Variability Selection Hypothesis allows for a more flexible interpretation of the evolution of bipedalism in hominins and a more fluid interpretation of the Turnover Pulse Hypothesis, where species turnover is meant to be more rapid. In some ways, this hypothesis accommodates both environmental data and our interpretations of an evolution toward greater variability among species and the survivability of generalists.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000;background-color: #ff99cc\"><strong>Paleoenvironment Summary<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff99cc\">Some hypotheses presented in this section pay specific attention to habitat (Savannah Hypothesis) while others point to large-scale climatic forces (Variability Selection Hypothesis). Some may be interpreted to describe the evolution of traits such as bipedalism (Savannah Hypothesis), and others generally explain the diversification of early hominins (Turnover Pulse and Variability Selection Hypotheses). While there is no consensus as to how the environment drove our evolution, it is clear that the environment shaped both habitat and resource availability in ways that would have influenced our early ancestors physically and behaviorally.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Derived Adaptations: Bipedalism<br \/>\n<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The unique form of locomotion exhibited by modern humans, called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1332\">obligate bipedalism<\/a><\/strong>, is important in distinguishing our species from the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1334\"><strong>extant<\/strong><\/a> (living) great apes. The ability to walk habitually upright is thus considered one of the defining attributes of the hominin lineage. We also differ from other animals that walk bipedally (such as kangaroos) in that we do not have a tail to balance us as we move.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The origin of bipedalism in hominins has been debated in paleoanthropology, but at present there are two main <span style=\"background-color: #ff99cc\">ideas<\/span>: <span style=\"text-decoration: underline\">(<em>theories<\/em>)<\/span><\/span><\/p>\n<ol>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">early hominins initially lived in trees, but increasingly started living on the ground, so we were a product of an arboreal last common ancestor (LCA) or,<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">our LCA was a terrestrial quadrupedal knuckle-walking species, more similar to extant chimpanzees.<\/span><\/li>\n<\/ol>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Most research supports the first theory of an arboreal LCA based on skeletal morphology of early hominin genera that demonstrate adaptations for climbing but not for knuckle-walking. This would mean that both humans and chimpanzees can be considered \u201cderived\u201d in terms of locomotion since chimpanzees would have independently evolved knuckle-walking.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are many current ideas regarding selective pressures that would lead to early hominins adapting upright posture and locomotion. Many of these selective pressures, as we have seen in the previous section, coincide with a shift in environmental conditions, supported by paleoenvironmental data. In general, however, it appears that, like extant great apes, early hominins thrived in forested regions with dense tree coverage, which would indicate an arboreal lifestyle. As the environmental conditions changed and a savannah\/grassland environment became more widespread, the tree cover would become less dense, scattered, and sparse such that bipedalism would become more important.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are several proposed selective pressures for bipedalism:<\/span><\/p>\n<ol>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\"><strong>E<\/strong><strong>nergy conservation:<\/strong> Modern bipedal humans conserve more energy than extant chimpanzees, which are predominantly knuckle-walking quadrupeds when walking over land. While chimpanzees, for instance, are faster than humans terrestrially, they expend large amounts of energy being so. Adaptations to bipedalism include \u201cstacking\u201d the majority of the weight of the body over a small area around the center of gravity (i.e., the head is above the chest, which is above the pelvis, which is over the knees, which are above the feet). This reduces the amount of muscle needed to be engaged during locomotion to \u201cpull us up\u201d and allows us to travel longer distances expending far less energy.<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\"><strong>T<\/strong><strong>hermoregulation:<\/strong> Less surface area (i.e., only the head and shoulders) is exposed to direct sunlight during the hottest parts of the day (i.e., midday). This means that the body has less need to employ additional \u201ccooling\u201d mechanisms such as sweating, which additionally means less water loss.<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Bipedalism <\/strong><span style=\"text-decoration: underline\">(Freeing of Hands)<\/span><strong>: <\/strong>This method of locomotion freed up our ancestors\u2019 hands such that they could more easily gather food and carry tools or infants. This further enabled the use of hands for more specialized adaptations associated with the manufacturing and use of tools.<\/span><\/li>\n<\/ol>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">These selective pressures are not mutually exclusive. Bipedality could have evolved from a combination of these selective pressures, in ways that increased the chances of early hominin survival.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Skeletal Adaptations for Bipedalism<\/strong><\/span><\/h3>\n<figure style=\"width: 405px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image10-1.png\" alt=\"A full human skeleton and gorilla skeleton standing in upright positions next to each other.\" width=\"405\" height=\"452\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.6: Compared to gorillas (right) and other apes, humans (left) have highly specialized adaptations to facilitate bipedal locomotion. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Primatenskelett-drawing.jpg\">Skeleton of human (1) and gorilla (2), unnaturally sketched<\/a> by unknown from Brehms Tierleben, Small Edition 1927 is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Humans have highly specialized adaptations to facilitate obligate bipedalism (Figure 9.6). Many of these adaptations occur within the soft tissue of the body (e.g., muscles and tendons). However, when analyzing the paleoanthropological record for evidence of the emergence of bipedalism, all that remains is the fossilized bone. Interpretations of locomotion are therefore often based on comparative analyses between fossil remains and the skeletons of extant primates with known locomotor behaviors. These adaptations occur throughout the skeleton and are summarized in Figure 9.7.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The majority of these adaptations occur in the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1336\"><strong>postcranium<\/strong><\/a> <span style=\"background-color: #ff99cc\">(the skeleton from below the head)<\/span> and are outlined in Figure 9.7. In general, these adaptations allow for greater stability and strength in the lower limb, by allowing for more shock absorption, for a larger surface area for muscle attachment, and for the \u201cstacking\u201d of the skeleton directly over the center of gravity to reduce energy needed to be kept upright. These adaptations often mean less flexibility in areas such as the knee and foot.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">However, these adaptations come at a cost. Evolving from a nonobligate bipedal ancestor means that the adaptations we have are evolutionary compromises. For instance, the valgus knee (angle at the knee) is an essential adaptation to balance the body weight above the ankle during bipedal locomotion. However, the strain and shock absorption at an angled knee eventually takes its toll. For example, runners often experience joint pain. Similarly, the long neck of the femur absorbs stress and accommodates for a larger pelvis, but it is a weak point, resulting in hip replacements being commonplace among the elderly, especially in cases where the bone additionally weakens through osteoporosis. Finally, the S-shaped curve in our spine allows us to stand upright, relative to the more curved C-shaped spine of an LCA. Yet the weaknesses in the curves can lead to pinching of nerves and back pain. Since many of these problems primarily are only seen in old age, they can potentially be seen as an evolutionary compromise.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Despite relatively few postcranial fragments, the fossil record in early hominins indicates a complex pattern of emergence of bipedalism. Key features, such as a more anteriorly placed foramen magnum, are argued to be seen even in the earliest discovered hominins, indicating an upright posture (Dart 1925). Some early species appear to have a mix of ancestral (arboreal) and derived (bipedal) traits, which indicates a mixed locomotion and a more <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1402\">mosaic evolution<\/a><\/strong> of the trait. Some early hominins appear to, for instance, have bowl-shaped pelvises (hip bones) and angled femurs suitable for bipedalism but also have retained an opposable <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1338\">hallux<\/a><\/strong> (big toe) or curved fingers and longer arms (for arboreal locomotion). These mixed morphologies may indicate that earlier hominins were not fully obligate bipeds and thus thrived in mosaic environments.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Yet the associations between postcranial and the more diagnostic cranial fossils and bones are not always clear, muddying our understanding of the specific species to which fossils belong (Grine et al. 2022).<\/span><\/p>\n<div align=\"left\">\n<table class=\"grid\">\n<caption>\n<p class=\"import-Normal\" style=\"text-align: left\"><span style=\"color: #000000\">Figure 9.7: Skeletal comparisons between modern humans (obligate bipeds) and nonobligate bipeds (e.g., chimpanzees). Credit: <a class=\"rId34\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\">Skeletal comparisons between modern humans and <\/a><a class=\"rId35\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\">nonobligate<\/a><a class=\"rId36\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\"> bipeds (Figure 9.6)<\/a> original to <a class=\"rId37\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a class=\"rId38\" style=\"color: #000000\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/span><\/p>\n<\/caption>\n<thead>\n<tr>\n<td style=\"width: 97.998px\"><strong>Region<\/strong><\/td>\n<td style=\"width: 106.992px\"><strong>Feature<\/strong><\/td>\n<td style=\"width: 366.992px\"><strong>Obligate Biped (H. sapiens)<\/strong><\/td>\n<td style=\"width: 310px\"><strong>Nonobligate Biped<\/strong><\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr>\n<td style=\"width: 97.998px\">Cranium<\/td>\n<td style=\"width: 106.992px\">Position of the foramen magnum<\/td>\n<td style=\"width: 366.992px\">Positioned inferiorly (immediately under the cranium) so that the head rests on top of the vertebral column for balance and support (head is perpendicular to the ground).<\/td>\n<td style=\"width: 310px\">Posteriorly positioned (to the back of the cranium). Head is positioned parallel to the ground.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Body proportions<\/td>\n<td style=\"width: 366.992px\">Shorter upper limb (not used for locomotion).<\/td>\n<td style=\"width: 310px\">Longer upper limbs (used for locomotion).<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Spinal curvature<\/td>\n<td style=\"width: 366.992px\">S-curve due to pressure exerted on the spine from bipedalism (lumbar lordosis).<\/td>\n<td style=\"width: 310px\">C-curve.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Vertebrae<\/td>\n<td style=\"width: 366.992px\">Robust lumbar (lower-back) vertebrae (for shock absorbance and weight bearing). Lower back is more flexible than that of apes as the hips and trunk swivel when walking (weight transmission).<\/td>\n<td style=\"width: 310px\">Gracile lumbar vertebrae compared to those of modern humans.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Pelvis<\/td>\n<td style=\"width: 366.992px\">Shorter, broader, bowl-shaped pelvis (for support); very robust. Broad sacrum with large sacroiliac joint surfaces.<\/td>\n<td style=\"width: 310px\">Longer, flatter, elongated ilia; more narrow and gracile; narrower sacrum; relatively smaller sacroiliac joint surface.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Lower limb<\/td>\n<td style=\"width: 366.992px\">In general, longer, more robust lower limbs and more stable, larger joints.<\/p>\n<ul>\n<li style=\"font-weight: 400\">Large femoral head and longer neck (absorbs more stress and increases the mechanical advantage).<\/li>\n<li style=\"font-weight: 400\">Valgus knee, in which the angle of the knee positions it over the ankle and keeps the center of gravity balanced over the stance leg during stride cycle (shock absorbance).<\/li>\n<li style=\"font-weight: 400\">Distal tibia (lower leg) of humans has a large medial malleolus for stability.<\/li>\n<\/ul>\n<\/td>\n<td style=\"width: 310px\">In general, smaller, more gracile limbs with more flexible joints.<\/p>\n<ul>\n<li style=\"font-weight: 400\">Femoral neck is smaller in comparison to modern humans and shorter.<\/li>\n<li style=\"font-weight: 400\">The legs bow outward, and there is no valgus angle of the knee (no \u201cknock knees\u201d).<\/li>\n<li style=\"font-weight: 400\">The distal tibia in chimpanzees is trapezoid (wider anteriorly) for climbing and allows more flexibility.<\/li>\n<\/ul>\n<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Foot<\/td>\n<td style=\"width: 366.992px\">Rigid, robust foot, without a midtarsal break.<\/p>\n<p>Nonopposable and large, robust big toe (for push off while walking) and large heel for shock absorbance.<\/td>\n<td style=\"width: 310px\">Flexible foot, midtarsal break present (which allows primates to lift their heels independently from their feet), opposable big toe for grasping.<\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<p class=\"import-Normal\"><span style=\"color: #000000;background-color: #ccffcc\">It is also worth noting that, while not directly related to bipedalism per se, other postcranial adaptations are evident in the hominin fossil record from some of the earlier hominins. For instance, the hand and finger morphologies of many of the earliest hominins indicate adaptations consistent with arboreality. These include longer hands, more curved metacarpals and phalanges (long bones in the hand and fingers, respectively), and a shorter, relatively weaker thumb. This allows for gripping onto curved surfaces during locomotion. The earliest hominins appear to have mixed morphologies for both bipedalism and arborealism. However, among Australopiths (members of the genus, Australopithecus), there are indications for greater reliance on bipedalism as the primary form of locomotion. Similarly, adaptations consistent with tool manufacture (shorter fingers and a longer, more robust thumb, in contrast to the features associated with arborealism) have been argued to appear before the genus <em>Homo<\/em>.<\/span><\/p>\n<div class=\"textbox shaded\">\n<h2>Special Topic: Fear of Snakes \u2014 A Cultural or Biological Response?<\/h2>\n<figure style=\"width: 245px\" class=\"wp-caption alignright\"><img src=\"https:\/\/www.researchgate.net\/profile\/David-Penning\/publication\/315305069\/figure\/fig2\/AS:613947958902785@1523388008174\/Constriction-coil-postures-of-kingsnakes-and-ratsnakes-Typical-constriction-coil.png\" alt=\"Constriction coil postures of kingsnakes and ratsnakes. Typical constriction coil postures in a kingsnake, Lampropeltis getula (92 g; A), and a ratsnake, Pantherophis guttatus (86 g; B). Both snakes were constricting similarly sized mice, Mus musculus (12 g). The relative prey mass was 13% for the kingsnake and 13.9% for the ratsnake.\u00a0\" width=\"245\" height=\"245\" \/><figcaption class=\"wp-caption-text\">Constriction coil postures of kingsnakes and ratsnakes. source: https:\/\/www.researchgate.net\/figure\/Constriction-coil-postures-of-kingsnakes-and-ratsnakes-Typical-constriction-coil_fig2_315305069<\/figcaption><\/figure>\n<p>It is suggested that primates have three major predators: raptors, felines, and snakes; however, many studies show that of these carnivores, snakes were one of the first that mammals had to contend with alongside dinosaurs, as felines and raptors evolved at a much slower pace than their reptilian competition. Herpetologists trace the evolution of constricting snakes to about 100 million years ago, and by the time mammals arrived around 75 million years ago, constrictors were\u00a0 already well established as a formidable threat (Greene, 2017). \u00a0Both co-existed for millennia and each sustained selective pressures requiring them to evolve specific traits to survive. When venomous snakes eventually emerged 55 to 65 million years ago, they posed yet an additional threat to proto-primates as they required less distance for the predator to kill (2017). Alongside camouflage and silent movement techniques, it was the development of the snake\u2019s hollow fangs through which to deliver venom that was most transformative to primate evolution. As such, primates evolved their pre-conscious attention, and visual acuity to cope with this new threat; therefore, while snakes were adapting morphologically to feed themselves, they were unwittingly teaching proto-primates valuable lessons in predator detection and reacting appropriately in order to survive.<\/p>\n<p>In a 2009 Harvard University study, Lynne A. Isbell hypothesizes that envenoming snakes are linked to being directly responsible for the origins of the evolving complex brains and superior visual capacity in the lineage of anthropoids leading to humans (Isbell, 2009). Forward-facing eyes for binocular vision, depth perception, enhanced visual acuity, stereoscopic and trichromatic colour vision, all traits necessary for snake detection; and the quick motor responses from the primate\u2019s fight, flight, or freeze defence mechanism to circumvent a snake\u2019s squeeze or bite. Numerous laboratory studies show that humans and primates both sense and visually detect snakes more rapidly than other threatening stimuli (Van Le Et al., 2013). These experiments show that snakes elicited the strongest, fastest responses (Van Le Et al., 2013). This is known as \u2018Snake Detection Theory\u2019 and is the evolution of the primate\u2019s complex brain, visual acuity, and rapid motor responses towards snakes in its environment that are the adaptations needed to live successfully as arboreal beings. It is not fortuitous then, that primates that never coexisted with venomous snakes, such as lemurs in Madagascar, have less visual acuity, better olfaction and smaller brains. Within Isbell\u2019s work, a collaborative study by a group of neuroscientists tested this hypothesis and found that, indeed, there is higher neural firing and activity in multiple areas of the primate brain, notably in the pulvinar, a region\u00a0 responsible for visual attention and oculomotor behaviour (Isbell, L., 2009).<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 509px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/www.worldhistory.org\/img\/r\/p\/750x750\/4941.jpg?v=1747098791\" alt=\"Apophis Defeated\" width=\"509\" height=\"309\" \/><figcaption class=\"wp-caption-text\">Tomb of Inherkau no. 359 Second chamber, South wall \"The great cat of Heliopolis\" killing the enemy of the sun, Apophis. source: https:\/\/www.worldhistory.org\/image\/4941\/apophis-defeated\/<\/figcaption><\/figure>\n<p>Today, the fear of snakes is widespread in humans, often shown through avoidance and disgust. A study in <em data-start=\"197\" data-end=\"244\">The Journal of Ethnobiology and Ethnomedicine<\/em> notes that snakes are over-hunted and excluded from conservation efforts worldwide (Cer\u00edaco, 2012). While cultural factors shape our sentiments, instinct also plays a role\u2014such as the developed avoidance behaviors toward threats like snakes. This blend of instinct and cultural influence is not only seen in behavior but also deeply embedded in the stories we tell. Many cultures depict mythological snakes as harbingers of death or chaos. In the Bible, Satan becomes a snake to tempt Eve. Norse mythology features J\u00f6rmungandr, the world serpent who signals the apocalypse. Egyptian myth tells of Apophis, who battles the sun god Ra nightly. Though sources vary, these myths consistently portray snakes as threats. As such, the widespread fear of snakes may reflect both evolutionary and cultural influences. Understood as an adaptive response inherited from primate ancestors\u2014who developed avoidance behaviors toward potentially dangerous stimuli\u2014and reinforced through myths and religious narratives, the enduring presence of snakes as potent figures of fear across human societies and primate groups highlights the complex intertwining of instinct and cultural meaning in shaping human behavior.<\/p>\n<\/div>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Early Hominins: Sahelanthropus and Orrorin<\/strong><\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">We see evidence for bipedalism in some of the earliest fossil hominins, dated from within our estimates of our divergence from chimpanzees. These hominins, however, also indicate evidence for arboreal locomotion.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The earliest dated hominin find (between 6 mya and 7 mya, based on radiometric dating of volcanic tufts) has been argued to come from Chad and is named <strong><em>Sahelanthropus tchadensis<\/em> <\/strong>(Figure 9.8; Brunet et al. 1995). The initial discovery was made in 2001 by Ahounta Djimdoumalbaye and announced in <em>Nature<\/em> in 2002 by a team led by French paleontologist Michel Brunet. The find has a small cranial capacity (360 cc) and smaller canines than those in extant great apes, though they are larger and pointier than those in humans. <span style=\"background-color: #ffff00\">This implies strongly that, over evolutionary time, the need for display and dominance among males has reduced, as has our sexual dimorphism<\/span>. A short cranial base and a foramen magnum that is more humanlike in positioning have been argued to indicate upright walking.<\/span><\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 640px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-289\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.82.jpg\" alt=\"Four views of a beige-colored skull are seen against a black background.\" width=\"640\" height=\"640\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.8: Sahelanthropus tchadensis exhibits a set of derived features, including a long, low cranium; a small, ape-sized braincase; and relatively reduced prognathism. Credit: aa <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 anterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 posterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 inferior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; and d <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 lateral left view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Initially, the inclusion of <em>Sahelanthropus<\/em> in the hominin family was debated by researchers, since the evidence for bipedalism is based on cranial evidence alone, which is not as convincing as postcranial evidence. Yet, a femur (thigh bone) and ulnae (upper arm bones) thought to belong to <em>Sahelanthropus<\/em> was discovered in 2001 (although not published until 2022). These bones may support the idea that the hominin was in fact a terrestrial biped with arboreal capabilities and behaviors (Daver et al. 2022).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Orrorin tugenensis<\/em><\/strong> (Orrorin meaning \u201coriginal man\u201d), dated to between 6 mya and 5.7 mya, was discovered near Tugen Hills in Kenya in 2000. Smaller <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1340\">cheek teeth<\/a><\/strong> (molars and premolars) than those in even more recent hominins, thick enamel, and reduced, but apelike, canines characterize this species. This is the first species that clearly indicates adaptations for bipedal locomotion, with fragmentary leg, arm, and finger bones having been found but few cranial remains. One of the most important elements discovered was a proximal femur, BAR 1002'00. The femur is the thigh bone, and the proximal part is that which articulates with the pelvis; this is very important for studying posture and locomotion. This femur indicates that <em>Ororrin<\/em> was bipedal, and recent studies suggest that it walked in a similar way to later <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1404\">Pliocene<\/a><\/strong> hominins. Some have argued that features of the finger bones suggest potential tool-making capabilities, although many researchers argue that these features are also consistent with climbing.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Early Hominins: The Genus <em>Ardipithecus<\/em><\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Another genus, <em>Ardipithecus<\/em>, is argued to be represented by at least two species: <em>Ardipithecus (Ar.) ramidus <\/em>and <em>Ar. kadabba<\/em>.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Ardipithecus ramidus<\/em><\/strong> (\u201cramid\u201d means root in the Afar language) is currently the best-known of the earliest hominins (Figure 9.9). Unlike <em>Sahelanthropus<\/em> and<em> Orrorin<\/em>, this species has a large sample size of over 110 specimens from Aramis alone. Dated to 4.4 mya, <em>Ar. ramidus<\/em> was found in Ethiopia (in the Middle Awash region and in Gona). This species was announced in 1994 by American palaeoanthropologist Tim White, based on a partial female skeleton nicknamed \u201cArdi\u201d (ARA-VP-6\/500; White et al. 1994). Ardi demonstrates a mosaic of ancestral and derived characteristics in the postcrania. For instance, she had an opposable big toe (hallux), similar to chimpanzees (i.e., more ancestral), which could have aided in climbing trees effectively. However, the pelvis and hip show that she could walk upright (i.e., it is derived), supporting her hominin status. A small brain (300 cc to 350 cc), midfacial projection, and slight prognathism show retained ancestral cranial features, but the cheek bones are less flared and robust than in later hominins.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 706px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-1310\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.9-scaled-1.jpg\" alt=\"Skull cast and partial skeleton with photographs of some bones and line drawings of others.\" width=\"706\" height=\"453\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.9a and b: Researchers believe that Ardipithecus ramidus was able to walk upright, although not as efficiently as later humans. It possessed the musculature required for tree climbing, and while moving quadrupedally, it likely placed weight on the palms of the hands rather than on the knuckles. Credit: a. <a class=\"rId61\" href=\"https:\/\/boneclones.com\/product\/ardipithecus-ramidus-skull-BH-039\">Ardipithecus ramidus Skull<\/a> by <a class=\"rId62\" href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a> is used by permission and available here under a <a class=\"rId63\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>; b. <a class=\"rId64\" href=\"https:\/\/boneclones.com\/product\/ardipithecus-ramidus-skull-BH-039\">Artist\u2019s rendition of \u201cArdi\u201d skeleton<\/a> by <a class=\"rId65\" href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a> is used by permission and available here under a <a class=\"rId66\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong><em>Ardipithecus kadabba<\/em><\/strong> (the species name means \u201coldest ancestor\u201d in the Afar language) is known from localities on the western margin of the Middle Awash region, the same locality where <em>Ar. ramidus<\/em> has been found. Specimens include mandibular fragments and isolated teeth as well as a few postcranial elements from the Asa Koma (5.5 mya to 5.77 mya) and Kuseralee Members (5.2 mya), well-dated and understood (but temporally separate) volcanic layers in East Africa. This species was discovered in 1997 by paleoanthropologist Dr. Yohannes Haile-Selassie. Originally these specimens were referred to as a subspecies of <em>Ar. ramidus<\/em>. In 2002, six teeth were discovered at Asa Koma and the dental-wear patterns confirmed that this was a distinct species, named <em>Ar. kadabba,<\/em> in 2004. One of the postcranial remains recovered included a 5.2 million-year-old toe bone that demonstrated features that are associated with toeing off (pushing off the ground with the big toe leaving last) during walking, a characteristic unique to bipedal walkers. However, the toe bone was found in the Kuseralee Member, and therefore some doubt has been cast by researchers about its association with the teeth from the Asa Koma Member.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"background-color: #ff99cc\"><strong><span style=\"color: #000000\">Bipedal Trends in Early Hominins: Summary<\/span><\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff99cc\">Trends toward bipedalism are seen in our earliest hominin finds. However, many specimens also indicate retained capabilities for climbing. Trends include a larger, more robust hallux; a more compact foot, with an arch; a robust, long femur, angled at the knee; a robust tibia; a bowl-shaped pelvis; and a more anterior foramen magnum. While the level of bipedality in <em>Salehanthropus<\/em> <em>tchadenisis<\/em> is debated since there are few fossils and no postcranial evidence, <em>Orrorin tugenensis<\/em> and <em>Ardipithecus<\/em> <em>k<\/em><em>adabba <\/em>show clear indications of some of these bipedal trends. However, some retained ancestral traits, such as an opposable hallux in <em>Ardipithecus<\/em>, indicate some retention in climbing ability.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Derived Adaptations: Early Hominin Dention<\/span><\/h2>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">The Importance of Teeth<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Teeth are abundant in the fossil record, primarily because they are already highly mineralized as they are forming, far more so than even bone. Because of this, teeth preserve readily. And, because they preserve readily, they are well-studied and better understood than many skeletal elements. In the sparse hominin (and primate) fossil record, teeth are, in some cases, all we have.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Teeth also reveal a lot about the individual from whom they came. We can tell what they evolved to eat, to which other species they may be closely related, and even, to some extent, the level of sexual dimorphism, or general variability, within a given species. This is powerful information that can be contained in a single tooth. With a little more observation, the wearing patterns on a tooth can tell us about the diet of the individual in the weeks leading up to its death. Furthermore, the way in which a tooth is formed, and the timing of formation, can reveal information about changes in diet (or even mobility) over infancy and childhood, using isotopic analyses. When it comes to our earliest hominin relatives, this information is vital for understanding how they lived.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The purpose of comparing different hominin species is to better understand the functional morphology as it applies to dentition. In this, we mean that the morphology of the teeth or masticatory system (which includes jaws) can reveal something about the way in which they were used and, therefore, the kinds of foods these hominins ate. When comparing the features of hominin groups, it is worth considering modern analogues (i.e., animals with which to compare) to make more appropriate assumptions about diet. In this way, hominin dentition is often compared with that of chimpanzees and gorillas (our close ape relatives), as well as with that of modern humans.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The most divergent group, however, is humans. Humans around the world have incredibly varied diets. Among hunter-gatherers, it can vary from a honey- and plant-rich diet, as seen in the Hadza in Tanzania, to a diet almost entirely reliant on animal fat and protein, as seen in Inuits in polar regions of the world. We are therefore considered generalists, more general than the largely <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1342\"><strong>frugivorous<\/strong><\/a> (fruit-eating) chimpanzee or the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1344\"><strong>folivorous<\/strong><\/a> (foliage-eating) gorilla, as discussed in Chapter 5.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">One way in which all humans are similar is our reliance on the processing of our food. We cut up and tear meat with tools using our hands, instead of using our front teeth (incisors and canines). We smash and grind up hard seeds, instead of crushing them with our hind teeth (molars). This means that, unlike our ape relatives, we can rely more on developing tools to navigate our complex and varied diets. <span style=\"text-decoration: underline\">(We could say)<\/span> Our brain, therefore, is our primary masticatory organ. Evolutionarily, our teeth have reduced in size and our faces are flatter, or more <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1346\">orthognathic<\/a>, <\/strong>partially in response to our increased reliance on our hands and brain to process food. Similarly, a reduction in teeth and a more generalist dental morphology could also indicate an increase in softer and more variable foods, such as the inclusion of more meat. <span style=\"background-color: #ffff00\">These trends begin early on in our evolution.<\/span> The link has been made between some of the earliest evidence for stone tool manufacture, the earliest members of our genus, and the features that we associate with these specimens.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">General Dental Trends in Early Hominins<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Several trends are visible in the dentition of early hominins. However, all tend to have the same <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1348\"><strong>dental formula<\/strong><\/a>. The dental formula tells us how many of each tooth type are present in each quadrant of the mouth. Going from the front of the mouth, this includes the square, flat <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1160\">incisors<\/a><\/strong>; the pointy <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1166\">canines<\/a><\/strong>; the small, flatter <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1350\"><strong>premolars<\/strong><\/a>; and the larger hind <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1164\">molars<\/a><\/strong>. In many primates, from Old World monkeys to great apes, the typical dental formula is 2:1:2:3. This means that if we divide the mouth into quadrants, each has two incisors, one canine, two premolars, and three molars. The eight teeth per quadrant total 32 teeth in all (although some humans have fewer teeth due to the absence of their wisdom teeth, or third molars).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 380px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image24.png\" alt=\"Anterior view of the lower face of a person showing their teeth.\" width=\"380\" height=\"253\" \/><figcaption class=\"wp-caption-text\">Figure 9.10: In humans, our canines are often a similar size to our incisors. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Adult_human_teeth.jpg\">Adult human teeth<\/a> by <a href=\"https:\/\/www.genusfotografen.se\/\">Genusfotografen<\/a> (Tomas Gunnarsson) through <a href=\"https:\/\/wikimedia.se\/\">Wikimedia Sverige<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p><span style=\"color: #000000\">The morphology of the individual teeth is where we see the most change. Among primates, large incisors are associated with food procurement or preparation (such as biting small fruits), while small incisors indicate a diet that may contain small seeds or leaves (where the preparation is primarily in the back of the mouth). Most hominins have relatively large, flat, vertically aligned incisors that <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1396\">occlude<\/a> <\/strong>(touch) relatively well, forming a \u201cbite.\u201d This differs from, for instance, the orangutan, whose teeth stick out (i.e.<em>,<\/em> are <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1352\"><strong>procumbent<\/strong><\/a>).<\/span><\/p>\n<p>&nbsp;<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">While the teeth are often aligned with diet, the canines may be misleading in that regard. We tend to associate pointy, large canines with the ripping required for meat, and the reduction (or, in some animals, the absence) of canines as indicative of herbivorous diets. In humans, our canines are often a similar size to our incisors and therefore considered <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1398\">incisiform<\/a><\/strong> (Figure 9.10). However, our closest relatives all have very long, pointy canines, particularly on their upper dentition. This is true even for the gorilla, which lives almost exclusively on plants. The canines in these instances reveal more about social structure and sexual dimorphism than diet, as large canines often signal <span style=\"background-color: #ffff00\">dominance<\/span>.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Early on in human evolution, we see a reduction in canine size. <em>Sahelanthropus tchadensis<\/em> and <em>Orrorin tugenensis<\/em> both have smaller canines than those in extant great apes, yet the canines are still larger and pointier than those in humans or more recent hominins. <span style=\"background-color: #ffff00\">This implies strongly that, over evolutionary time, the need for display and dominance among males has reduced, as has our sexual dimorphism.<\/span> In <em>Ardipithecus ramidus<\/em>, there is no obvious difference between male and female canine size, yet they are still slightly larger and pointier than in modern humans. This implies a less sexually dimorphic social structure in the earlier hominins relative to modern-day chimpanzees and gorillas.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Along with a reduction in canine size is the reduction or elimination of a canine <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1172\">diastema<\/a>:<\/strong> a gap between the teeth on the mandible that allows room for elongated teeth on the maxilla to \u201cfit\u201d in the mouth. Absence of a diastema is an excellent indication of a reduction in canine size. In animals with large canines (such as baboons), there is also often a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1354\"><strong>honing P3<\/strong><\/a>, where the first premolar (also known as P3 for evolutionary reasons) is triangular in shape, \u201csharpened\u201d by the extended canine from the upper dentition. This is also seen in some early hominins: <em>Ardipithecus<\/em>, for example, has small canines that are almost the same height as its incisors, although still larger than those in recent hominins.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The hind dentition, such as the bicuspid (two cusped) premolars or the much larger molars, are also highly indicative of a generalist diet in hominins. Among the earliest hominins, the molars are larger than we see in our genus, increasing in size to the back of the mouth and angled in such a way from the much smaller anterior dentition as to give these hominins a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1356\"><strong>parabolic<\/strong><\/a> (V-shaped) dental arch. This differs from our living relatives and some early hominins, such as <em>Sahelanthropus<\/em>, whose molars and premolars are relatively parallel between the left and right sides of the mouth, creating a U-shape.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Among more recent early hominins, the molars are larger than those in the earliest hominins and far larger than those in our own genus, <em>Homo.<\/em> Large, short molars with thick <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1358\"><strong>enamel<\/strong><\/a> allowed our early cousins to grind fibrous, coarse foods, such as sedges, which require plenty of chewing. This is further evidenced in the low <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1176\">cusps<\/a>,<\/strong> or ridges, on the teeth, which are ideal for chewing. In our genus, the hind dentition is far smaller than in these early hominins. Our teeth also have medium-size cusps, which allow for both efficient grinding and tearing\/shearing meats.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Understanding the dental morphology has allowed researchers to extrapolate very specific behaviors of early hominins. It is worth noting that while teeth preserve well and are abundant, a slew of other morphological traits additionally provide evidence for many of these hypotheses. Yet there are some traits that are ambiguous. For instance, while there are definitely high levels of sexual dimorphism in <em>Au. afarensis<\/em>, discussed in the next section, the canine teeth are reduced in size, implying that while canines may be useful indicators for sexual dimorphism, it is also worth considering other evidence.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff99cc\">In summary, trends among early hominins include a reduction in procumbency, reduced hind dentition (molars and premolars), a reduction in canine size (more incisiform with a lack of canine diastema and honing P3), flatter molar cusps, and thicker dental enamel. All early hominins have the ancestral dental formula of 2:1:2:3. These trends are all consistent with a generalist diet, incorporating more fibrous foods.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Special Topic: Contested Species<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Many named species are highly debated and argued to have specimens associated with a more variable <em>Au. afarensis <\/em>or <em>Au. anamensis<\/em> species. Sometimes these specimens are dated to times when, or found in places in which, there are \u201cgaps\u201d in the palaeoanthropological record. These are argued to represent chronospecies or variants of <em>Au. afarensis<\/em>. However, it is possible that, with more discoveries, the distinct species types will hold.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><strong><em>Australopithecus bahrelghazali<\/em><\/strong> is dated to within the time period of <em>Au. afarensi<\/em>s (3.6 mya; Brunet et al. 1995) and was the first Australopithecine to be discovered in Chad in central Africa. Researchers argue that the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1360\">holotype<\/a><\/strong>, whom discoverers have named \u201cAbel,\u201d falls under the range of variation of <em>Au. afarensis<\/em> and therefore that <em>A. bahrelghazali<\/em> does not fall into a new species (Lebatard et al. 2008). If \u201cAbel\u201d is a member of <em>Au. afarensis<\/em>, the geographic range of the species would be greatly extended.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">On a different note, <strong><em>Australopithecus <\/em><\/strong><strong><em>deyiremada<\/em><\/strong> (meaning \u201cclose relative\u201d in the Ethiopian language of Afar) is dated to 3.5 mya to 3.3 mya and is based on fossil mandible bones discovered in 2011 in Woranso-Mille (in the Afar region of Ethiopia) by Yohannes Haile-Selassie, an Ethiopian paleoanthropologist (Haile-Selassie et al. 2019). The discovery indicated, in contrast to <em>Au. afarensis<\/em>, smaller teeth with thicker enamel (potentially suggesting a harder diet) as well as a larger mandible and more projecting cheekbones. This find may be evidence that more than one closely related hominin species occupied the same region at the same temporal period (Haile-Selassie et al. 2015; Spoor 2015) or that other <em>Au. afarensis<\/em> specimens have been incorrectly designated. However, others have argued that this species has been prematurely identified and that more evidence is needed before splitting the taxa, since the variation appears subtle and may be due to slightly different niche occupations between populations over time.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><strong><em>Australopithecus garhi<\/em><\/strong> is another species found in the Middle Awash region of Ethiopia. It is currently dated to 2.5 mya (younger than <em>Au. afarensis<\/em>). Researchers have suggested it fills in a much-needed temporal \u201cgap\u201d between hominin finds in the region, with some anatomical differences, such as a relatively large cranial capacity (450 cc) and larger hind dentition than seen in other gracile Australopithecines. Similarly, the species has been argued to have longer hind limbs than <em>Au. afarensis<\/em>, although it was still able to move arboreally (Asfaw et al. 1999). However, this species is not well documented or understood and is based on only several fossil specimens. More astonishingly, crude stone tools resembling Oldowan (which will be described later) have been found in association with <em>Au. garhi<\/em>. While lacking some of the features of the Oldowan, this is one of the earliest technologies found in direct association with a hominin.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><strong><em>Kenyanthopus<\/em><\/strong><strong><em> platyops<\/em><\/strong> (the name \u201cplatyops\u201d refers to its flatter-faced appearance) is a highly contested genus\/species designation of a specimen (KNM-WT 40000) from Lake Turkana in Kenya, discovered by Maeve Leakey in 1999 (Figure 9.11). Dated to between 3.5 mya and 3.2 mya, some have suggested this specimen is an <em>Australopithecus<\/em>, perhaps even <em>Au.<\/em> <em>afarensis<\/em> (with a brain size which is difficult to determine, yet appears small), while still others have placed this specimen in <em>Homo <\/em>(small dentition and flat-orthognathic face). While taxonomic placing of this species is quite divided, the discoverers have argued that this species is ancestral to <em>Homo<\/em>, in particular to <em>Homo <\/em><em>ruldolfensis<\/em> (Leakey et al. 2001). Some researchers have additionally associated the earliest tool finds from Lomekwi, Kenya, temporally (3.3 mya) and in close geographic proximity to this specimen.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 579px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-292 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.11.jpg\" alt=\"Four views of an ancient skull are shown on a black background.\" width=\"579\" height=\"579\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.11: This specimen, KNM WT 40000 (Kenyanthopus platyops), has small detention, a small brain case, and a relatively flat face. Its genus\/species designation remains contested. Credit: a. <a class=\"rId76\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId77\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 anterior view<\/a> by \u00a9<a class=\"rId78\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId79\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId80\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId81\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId82\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 superior view<\/a> by \u00a9<a class=\"rId83\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId84\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId85\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a class=\"rId86\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId87\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 lateral left view<\/a> by \u00a9<a class=\"rId88\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId89\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId90\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; d. <a class=\"rId91\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId92\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 inferior view<\/a> by \u00a9<a class=\"rId93\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId94\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId95\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">The Genus <em>Australopithecus<\/em><br \/>\n<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The Australopithecines are a diverse group of hominins, comprising various species. <em>Australopithecus<\/em> is the given group or genus name. It stems from the Latin word <em>Australo<\/em>, meaning \u201csouthern,\u201d and the Greek word <em>pithecus,<\/em> meaning \u201cape.\u201d Within this section, we will outline these differing species\u2019 geological and temporal distributions across Africa, unique derived and\/or shared traits, and importance in the fossil record.<\/span><\/p>\n<figure style=\"width: 381px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image37-2.png\" alt=\"A skull has a pronounced sagittal crest, flaring cheekbones, and large hind teeth.\" width=\"381\" height=\"585\" \/><figcaption class=\"wp-caption-text\">Figure 9.12: Robust Australopithecines such as Paranthropus boisei had large molars and chewing muscles. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Paranthropus_boisei_skull.jpg\">Paranthropus boisei skull<\/a> by Durova is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/deed.en\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Between 3 mya and 1 mya, there seems to be differences in dietary strategy between different species of hominins designated as Australopithecines. A pattern of larger posterior dentition (even relative to the incisors and canines in the front of the mouth), thick enamel, and cranial evidence for extremely large chewing muscles is far more pronounced in a group known as the robust australopithecines. This pattern is extreme<span style=\"text-decoration: underline\">ly<\/span> relative to their earlier contemporaries or predecessors, the gracile australopithecines<strong>,<\/strong> and is certainly larger than those seen in early <em>Homo<\/em>, which emerged during this time. This pattern of incredibly large hind dentition (and very small anterior dentition) has led people to refer to robust australopithecines as <strong>megadont<\/strong> hominins (Figure 9.12).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Because of these differences, this section has been divided into \u201cgracile\u201d and \u201crobust\u201d Australopithecines, highlighting the morphological differences between the two groups (which many researchers have designated as separate genera: <em>Australopithecus<\/em> and <em>Paranthropus<\/em>, respectively) and then focusing on the individual species. It is worth noting, however, that not all researchers accept these clades as biologically or genetically distinct, with some researchers insisting that the relative gracile and robust features found in these species are due to parallel evolutionary events toward similar dietary niches.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Despite this genus\u2019 ancestral traits and small cranial capacity, all members show evidence of bipedal locomotion. It is generally accepted that <em>Australopithecus <\/em>species display varying degrees of arborealism along with bipedality.<\/span><\/p>\n<h3 class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Gracile Australopithecines<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">This section describes individual species from across Africa. These species are called \u201c<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1406\">gracile<\/a> <\/strong>australopithecines\u201d because of their smaller and less robust features compared to the divergent \u201c<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1408\"><strong>robust<\/strong><\/a>\u201d group. Numerous Australopithecine species have been named, but some are only based on a handful of fossil finds, whose designations are controversial.<\/span><\/p>\n<h4 class=\"import-Normal\"><em><span style=\"color: #000000\">East African Australopithecines<\/span><\/em><\/h4>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">East African Australopithecines are found throughout the EARS, and they include the earliest species associated with this genus. Numerous fossil-yielding sites, such as Olduvai, Turkana, and Laetoli, have excellent, datable stratigraphy, owing to the layers of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1362\"><strong>volcanic tufts<\/strong> <\/a> that have accumulated over millions of years. <span style=\"background-color: #ff9900\">These tufts may be dated using absolute dating techniques, such as Potassium-Argon dating (described in Chapter 7). This means that it is possible to know a relatively refined date for any fossil if the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1364\"><strong>context<\/strong> <\/a> (i.e., exact location) of that find is known.<\/span> Similarly, comparisons between the faunal assemblages of these stratigraphic layers have allowed researchers to chronologically identify environmental changes.<\/span><\/p>\n<figure style=\"width: 313px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image20-1-1.png\" alt=\"Occlusal view of an Au. anamensis mandible, with relatively large teeth, including canines.\" width=\"313\" height=\"313\" \/><figcaption class=\"wp-caption-text\">Figure 9.13: As seen in this mandible of KNM-KP 29281, Australopithecus anamensis had relatively large canine teeth. Credit: <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20anamensis\/KNM-KP%2029281\">Australopithecus anamensis: KNM-KP 29281 occlusal view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p><span style=\"color: #000000\">The earliest known Australopithecine is dated to 4.2 mya to 3.8 mya. <strong><em>Australopithecus anamensis<\/em><\/strong> (after \u201cAnam,\u201d meaning \u201clake\u201d from the Turkana region in Kenya; Leakey et al. 1995; Patterson and Howells 1967) is currently found from sites in the Turkana region (Kenya) and Middle Awash (Ethiopia; Figure 9.13). Recently, a 2019 find from Ethiopia, named MRD, after Miro Dora where it was found, was discovered by an Ethiopian herder named Ali Bereino. It is one of the most complete cranial finds of this species (Ward et al. 1999). A small brain size (370 cc), relatively large canines, projecting cheekbones, and earholes show more ancestral features as compared to those of more recent Australopithecines. The most important element discovered with this species is a fragment of a tibia (shinbone), which demonstrates features associated with weight transfer during bipedal walking. Similarly, the earliest found hominin femur belongs to this species. Ancestral traits in the upper limb (such as the humerus) indicate some retained arboreal locomotion.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Some researchers suggest that <em>Au. anamensis<\/em> is an intermediate form of the chronospecies that becomes <em>Au. afarensis<\/em>, evolving from <em>Ar. ramidus<\/em>. However, this is debated, with other researchers suggesting morphological similarities and affinities with more recent species instead. Almost 100 specimens, representing over 20 individuals, have been found to date (Leakey et al. 1995; McHenry 2009; Ward et al. 1999).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Au. afarensis<\/em><\/strong> is one of the oldest and most well-known australopithecine species and consists of a large number of fossil remains. <em>Au. afarensis<\/em> (which means \u201cfrom the Afar region\u201d) is dated to between 2.9 mya and 3.9 mya and is found in sites all along the EARS system, in Tanzania, Kenya, and Ethiopia (Figure 9.14). The most famous individual from this species is a partial female skeleton discovered in Hadar (Ethiopia), later nicknamed \u201cLucy,\u201d after the Beatles\u2019 song \u201cLucy in the Sky with Diamonds,\u201d which was played in celebration of the find (Johanson et al. 1978; Kimbel and Delezene 2009). This skeleton was found in 1974 by Donald Johanson and dates to approximately 3.2 mya. In addition, in 2002 a juvenile of the species was found by Zeresenay Alemseged and given the name \u201cSelam\u201d (meaning \u201cpeace,\u201d DIK 1-1), though it is popularly known as \u201cLucy\u2019s Child\u201d or as the \u201cDikika Child\u201d (Alemseged et al. 2006). Similarly, the \u201cLaetoli Footprints\u201d (discussed in Chapter 7; Hay and Leakey 1982; Leakey and Hay 1979) have drawn much attention.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 643px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-295 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.14.jpg\" alt=\"Two images of life-like reconstructions of female and male Au. afarensis.\" width=\"643\" height=\"322\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.14 a-b: Artistic reconstructions of Australopithecus afarensis by artist John Gurche. Female \u201cLucy\u201d is left and a male is on the right. Credit: a. <a class=\"rId106\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Australopithecus afarensis, \u201cLucy,\u201d adult female. Reconstruction based on AL-288-1 by artist John Gurche, front view close-up<\/a> by <a class=\"rId107\" href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?\u201d] is <a class=\"rId108\" href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>; b. <a class=\"rId109\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Australopithecus afarensis, adult male. Reconstruction based on <\/a><a class=\"rId110\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">AL444-2<\/a><a class=\"rId111\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\"> by John Gurche<\/a> by <a class=\"rId112\" href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?\u201d] is <a class=\"rId113\" href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>.<\/figcaption><\/figure>\n<figure style=\"width: 320px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image40.png\" alt=\"A partial skeleton includes bones of the cranium, mandible, and postcranium.\" width=\"320\" height=\"772\" \/><figcaption class=\"wp-caption-text\">Figure 9.15: The humanlike femoral neck, valgus knee, and bowl-shaped hip seen in the \u201cLucy\u201d skeleton indicates that Australopithecus afarensis was bipedal. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Lucy_blackbg.jpg\">Lucy blackbg<\/a> [AL 288-1, Australopithecus afarensis, cast from Museum national d'histoire naturelle, Paris] by 120 is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The canines and molars of <em>Au. afarensis<\/em> are reduced relative to great apes but are larger than those found in modern humans (indicative of a generalist diet); in addition, <em>Au. afarensis <\/em>has a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1366\"><strong>prognathic<\/strong> <\/a> face (the face below the eyes juts anteriorly) and robust facial features that indicate relatively strong chewing musculature (compared with <em>Homo<\/em>) but which are less extreme than in <em>Paranthropus<\/em>. Despite a reduction in canine size in this species, large overall size variation indicates high levels of sexual dimorphism.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Skeletal evidence indicates that this species was bipedal, as its pelvis and lower limb demonstrate a humanlike femoral neck, valgus knee, and bowl-shaped hip (Figure 9.15). <span style=\"background-color: #ccffcc\">More evidence of bipedalism is found in the footprints of this species<\/span>. <em>Au. afarensis<\/em> is associated with the Laetoli Footprints, <span style=\"background-color: #ff9900\">a 24-meter trackway of hominin fossil footprints preserved in volcanic ash discovered by Mary Leakey in Tanzania and dated to 3.5 mya to 3 mya. This set of prints is thought to have been produced by three bipedal individuals as there are no knuckle imprints, no opposable big toes, and a clear arch is present. The infants of this species are thought to have been more arboreal than the adults, as discovered through analyses of the foot bones of the Dikika Child dated to 3.32 mya (Alemseged et al. 2006).<\/span><\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Although not found in direct association with stone tools, potential evidence for cut marks on bones, found at Dikika, and dated to 3.39 mya indicates a possible temporal\/ geographic overlap between meat eating, tool use, and this species. However, this evidence is fiercely debated. Others have associated the cut marks with the earliest tool finds from Lomekwi, Kenya, temporally (3.3 mya) and in close geographic proximity to this species.<\/span><\/p>\n<h4 class=\"import-Normal\"><em><span style=\"color: #000000\">South African Australopithecines<\/span><\/em><\/h4>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Since the discovery of the Taung Child, there have been numerous Australopithecine discoveries from the region known as \u201cThe Cradle of Humankind,\u201d which was recently given UNESCO World Heritage Site status as \u201cThe Fossil Hominid Sites of South Africa.\u201d The limestone caves found in the Cradle allow for the excellent preservation of fossils. Past animals navigating the landscape and falling into cave openings, or caves used as dens by carnivores, led to the accumulation of deposits over millions of years. Many of the hominin fossils, encased in <strong>breccia<\/strong> (hard, calcareous sedimentary rock), are recently exposed from limestone quarries mined in the previous century. This means that extracting fossils requires excellent and detailed exposed work, often by a team of skilled technicians.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">While these sites have historically been difficult to date, with mixed assemblages accumulated over large time periods, advances in techniques such as uranium-series dating have allowed for greater accuracy. Historically, the excellent faunal record from East Africa has been used to compare sites based on <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1368\"><strong>relative dating<\/strong>,<\/a> whereby environmental and faunal changes and extinction events allow us to know which hominin finds are relatively younger or older than others.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The discovery of the Taung Child in 1924 (discussed in the Special Topic box \u201cThe Taung Child\u201d below) shifted the focus of palaeoanthropological research from Europe to Africa, although acceptance of this shift was slow (Broom 1947; Dart 1925). The species to which it is assigned, <strong><em>Australopithecus africanus<\/em><\/strong> (name meaning \u201cSouthern Ape of Africa\u201d), is currently dated to between 3.3 mya and 2.1 mya (Pickering and Kramers 2010), with discoveries from Sterkfontein, Taung, Makapansgat, and Gladysvale in South Africa (Figure 9.16). A relatively large brain (400 cc to 500 cc), small canines without an associated diastema, and more rounded cranium and smaller teeth than <em>Au. afarensis<\/em> indicate some derived traits. Similarly, the postcranial remains (in particular, the pelvis) indicate bipedalism. However, the sloping face and curved phalanges (indicative of retained arboreal locomotor abilities) show some ancestral features. Although not in direct association with stone tools, a 2015 study noted that the trabecular bone morphology of the hand was consistent with forceful tool manufacture and use, suggesting potential early tool abilities.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 570px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image19-1.png\" alt=\"A life-like reconstruction of the face of Australopithecus africanus, smiling in anterior view.\" width=\"570\" height=\"570\" \/><figcaption class=\"wp-caption-text\">Figure 9.16: An artistic reconstruction of Australopithecus africanus by John Gurche. Credit: <a href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Australopithecus africanus. Reconstruction based on STS 5 by John Gurche <\/a>by <a href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?] is <a href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Another famous <em>Au. africanus<\/em> skull (the skull of \u201cMrs. Ples\u201d) was previously attributed to <em>Plesianthropus transvaalensis<\/em><em>, <\/em>meaning \u201cnear human from the Transvaal,\u201d the old name for Gauteng Province, South Africa (Broom 1947, 1950). The name was shortened by contemporary journalists to \u201cPles\u201d (Figure 9.17). Due to the prevailing mores of the time, the assumed female found herself married, at least in name, and has become widely known as \u201cMrs. Ples.\u201d It was later reassigned to <em>Au. africanus<\/em> and is now argued by some to be a young male rather than an adult female cranium (Thackeray 2000, Thackeray et al. 2002).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 548px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-298 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.17.jpg\" alt=\"Four views of an ancient skull are shown on a black background.\" width=\"548\" height=\"548\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.17: The \u201cMrs. Ples\u201d brain case is small in size (like apes) but its face is less prognathic; its foramen magnum is positioned more like a modern human than an African apes. Credit: a. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 anterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 posterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 superior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; and d. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 lateral right view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In 2008, nine-year-old Matthew Berger, son of paleoanthropologist Lee Berger, noted a clavicle bone in some leftover mining breccia in the Malapa Fossil Site (South Africa). After rigorous studies, the species, <strong><em>Australopithecus sediba<\/em><\/strong> (meaning \u201cfountain\u201d or \u201cwellspring\u201d in the South African language of Sesotho), was named in 2010 (Figure 9.18; Berger et al. 2010). The first type specimen belongs to a juvenile male, Karabo (MH1), but the species is known from at least six partial skeletons, from infants through adults. These specimens are currently dated to 1.97 mya (Dirks et al. 2010). The discoverers have argued that <em>Au. sediba<\/em> shows mosaic features between <em>Au. africanus<\/em> and the genus, <em>Homo<\/em>, which potentially indicates a transitional species, although this is heavily debated. These features include a small brain size (<em>Australopithecus<\/em>-like; 420 cc to 450 cc) but gracile mandible and small teeth (<em>Homo<\/em>-like). Similarly, the postcranial skeletons are also said to have mosaic features: scientists have interpreted this mixture of traits (such as a robust ankle but evidence for an arch in the foot) as a transitional phase between a body previously adapted to arborealism (particularly in evidence from the bones of the wrist) to one that adapted to bipedal ground walking. Some researchers have argued that <em>Au. sediba<\/em> shows a modern hand morphology (shorter fingers and a longer thumb), indicating that adaptations to tool manufacture and use may be present in this species.<\/span><\/p>\n<figure style=\"width: 531px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image17-1.png\" alt=\"A beige-colored skull with no mandible on a black background has some missing teeth.\" width=\"531\" height=\"400\" \/><figcaption class=\"wp-caption-text\">Figure 9.18: Australopithecus sediba shows mosaic features between Au. africanus and Homo. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Australopithecus_sediba.JPG\">Australopithecus sediba<\/a>, photo by Brett Eloff courtesy <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Profberger\">Profberger<\/a> and <a href=\"https:\/\/en.wikipedia.org\/wiki\/University_of_the_Witwatersrand\">Wits University<\/a>, is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Another famous Australopithecine find from South Africa is that of the nearly complete skeleton now known as \u201cLittle Foot\u201d (Clarke 1998, 2013). Little Foot (StW 573) is potentially the earliest dated South African hominin fossil, dating to 3.7 mya, based on radiostopic techniques, although some argue that it is younger than 3 mya (Pickering and Kramers 2010). The name is jokingly in contrast to the cryptid species \u201cbigfoot\u201d and is named because the initial discovery of four ankle bones indicated bipedality. Little Foot was discovered by Ron Clarke in 1994, when he came across the ankle bones while sorting through monkey fossils in the University of Witwatersrand collections (Clarke and Tobias 1995). He asked Stephen Motsumi and Nkwane Molefe to identify the known records of the fossils, which allowed them to find the rest of the specimen within just days of searching the Sterkfontein Caves\u2019 Silberberg Grotto.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The discoverers of Little Foot insist that other fossil finds, previously identified as <em>Au. Africanus<\/em>, be placed in this new species based on shared ancestral traits with older East African Australopithecines (Clarke and Kuman 2019). These include features such as a relatively large brain size (408 cc), robust zygomatic arch, and a flatter midface. Furthermore, the discoverers have argued that the heavy anterior dental wear patterns, relatively large anterior dentition, and smaller hind dentition of this specimen more closely resemble that of <em>Au. anamensis<\/em> or <em>Au. afarensis<\/em>. It has thus been placed in the species <strong><em>Australopithecus prometheus<\/em><\/strong>. This species name refers to a previously defunct taxon named by Raymond Dart. The species designation was, through analyzing Little Foot, revived by Ron Clarke, who insists that many other fossil hominin specimens have prematurely been placed into <em>Au. africanus<\/em>. Others say that it is more likely that <em>Au. africanus<\/em> is a more variable species and not representative of two distinct species.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\"><em>Paranthropus<\/em> \u201cRobust\u201d Australopithecines<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In the robust australopithecines, the specialized nature of the teeth and masticatory system, such as flaring zygomatic arches (cheekbones), accommodate very large temporalis (chewing) muscles. These features also include a large, broad, dish-shaped face and and a large mandible with extremely large posterior dentition (referred to as megadonts) and hyper-thick enamel (Kimbel 2015; Lee-Thorp 2011; Wood 2010). Research has revolved around the shared adaptations of these \u201crobust\u201d australopithecines, linking their morphologies to a diet of hard and\/or tough foods (Brain 1967; Rak 1988). Some argued that the diet of the robust australopithecines was so specific that any change in environment would have accelerated their extinction. The generalist nature of the teeth of the gracile australopithecines, and of early <em>Homo<\/em>, would have made them more capable of adapting to environmental change. However, some have suggested that the features of the robust australopithecines might have developed as an effective response to what are known as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1370\"><strong>fallback <\/strong><strong>foods<\/strong><\/a> in hard times rather than indicating a lack of adaptability.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are currently three widely accepted robust australopithecus or, <em>Paranthropus<\/em>, species: <em>P. aethiopic<\/em><em>us<\/em>, which has more ancestral traits, and <em>P. boisei and P. robustus<\/em>, which are more derived in their features (Strait et al. 1997; Wood and Schroer 2017). These three species have been grouped together by a majority of scholars as a single genus as they share more derived features (are more closely related to each other; or, in other words, are <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1372\"><strong>monophyletic<\/strong><\/a>) than the other australopithecines (Grine 1988; Hlazo 2015; Strait et al. 1997; Wood 2010 ). While researchers have mostly agreed to use the umbrella term <em>Paranthropus<\/em>, there are those who disagree (Constantino and Wood 2004, 2007; Wood 2010).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">As a collective, this genus spans 2.7 mya to 1.0 mya, although the dates of the individual species differ. The earliest of the Paranthropus species, <strong><em>Paranthropus aethiopicus<\/em><\/strong>, is dated to between 2.7 mya and 2.3 mya and currently found in Tanzania, Kenya, and Ethiopia in the EARS system (Figure 9.19; Constantino and Wood 2007; Hlazo 2015; Kimbel 2015; Walker et al. 1986; White 1988). It is well known because of one specimen known as the \u201cBlack Skull\u201d (KNM\u2013WT 17000), so called because of the mineral manganese that stained it black during fossilization (Kimbel 2015). As with all robust Australopithecines, <em>P. aethiopicus<\/em> has the shared derived traits of large, flat premolars and molars; large, flaring zygomatic arches for accommodating large chewing muscles (the temporalis muscle); a sagittal crest (ridge on the top of the skull) for increased muscle attachment of the chewing muscles to the skull; and a robust mandible and supraorbital torus (brow ridge). However, only a few teeth have been found. A proximal tibia indicates bipedality and similar body size to <em>Au. afarensis<\/em>. In recent years, researchers have discovered and assigned a proximal tibia and juvenile cranium (L.338y-6) to the species (Wood and Boyle 2016).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 666px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-300 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.19.jpg\" alt=\"Five views of a beige partial skull on a black background.\" width=\"666\" height=\"444\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.19: The \u201cBlack Skull\u201d (Paranthropus aethiopicus) had a large sagittal crest and large, flared zygomatic arches that indicate it had large chewing muscles and a powerful biting force. Credit: a. <a class=\"rId156\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId157\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 anterior view<\/a> by \u00a9<a class=\"rId158\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId159\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId160\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId161\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId162\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 lateral right view<\/a> by \u00a9<a class=\"rId163\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId164\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId165\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a class=\"rId166\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId167\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 superior view<\/a> by \u00a9<a class=\"rId168\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId169\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId170\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; d. <a class=\"rId171\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId172\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 posterior view<\/a> by \u00a9<a class=\"rId173\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId174\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId175\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; e. <a class=\"rId176\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId177\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 inferior view<\/a> by \u00a9<a class=\"rId178\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId179\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId180\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">First attributed as <em>Zinjanthropus boisei<\/em> (with the first discovery going by the nickname \u201cZinj\u201d or sometimes \u201cNutcracker Man\u201d), <strong><em>Paranthropus boisei<\/em><\/strong> was discovered in 1959 by Mary Leakey (see Figure 9.20 and 9.21; Hay 1990; Leakey 1959). This \u201crobust\u201d australopith species is distributed across countries in East Africa at sites such as Kenya (Koobi Fora, West Turkana, and Chesowanja), Malawi (Malema-Chiwondo), Tanzania (Olduvai Gorge and Peninj), and Ethiopia (Omo River Basin and Konso). The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1374\">hypodigm<\/a><\/strong>, sample of fossils whose features define the group, has been found by researchers to date to roughly 2.4 mya to 1.4 mya. Due to the nature of its exaggerated, larger, and more robust features, <em>P. boisei <\/em>has been termed <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1376\"><strong>hyper-robust<\/strong><\/a>\u2014that is, even more heavily built than other robust species, with very large, flat posterior dentition (Kimbel 2015). Tools dated to 2.5 mya in Ethiopia have been argued to possibly belong to this species. Despite the cranial features of <em>P. boisei<\/em> indicating a tough diet of tubers, nuts, and seeds, isotopes indicate a diet high in C4 foods (e.g., grasses, such as sedges). Another famous specimen from this species is the Peninj mandible from Tanzania, found in 1964 by Kimoya Kimeu.<\/span><\/p>\n<figure style=\"width: 557px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image14-1.png\" alt=\"Life-like reconstruction of Paranthropus boisei.\" width=\"557\" height=\"557\" \/><figcaption class=\"wp-caption-text\">Figure 9.20: Artistic reconstruction of a Paranthropus boisei, male, by John Gurche. Credit: <a href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Paranthropus boisei, male. Reconstruction based on OH 5 and KNM-ER 406 by John Gurche<\/a> by <a href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?\u201d] is <a href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>.<\/figcaption><\/figure>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 565px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-302 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.21.jpg\" alt=\"Three views of an ancient skull are shown on a black background.\" width=\"565\" height=\"565\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.21: \u201cNutcracker Man\u201d (Paranthropus boisei) had hyper-robust features including very large dentition, flaring zygomatic arches, a broad concave face. It had a powerful and extremely efficient chewing force. Credit: <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20boisei\/OH%205\">Paranthropus boisei: OH 5 anterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20boisei\/OH%205\">Paranthropus boisei: OH 5 inferior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20boisei\/OH%205\">Paranthropus boisei: OH 5 posterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Paranthropus robustus<\/em><\/strong> was the first taxon to be discovered within the genus in Kromdraai B by a schoolboy named Gert Terblanche; subsequent fossil discoveries were made by researcher Robert Broom in 1938 (Figure 9.22; Broom 1938a, 1938b, 1950), with the holotype specimen TM 1517 (Broom 1938a, 1938b, 1950; Hlazo 2018). <em>Paranthropus robustus<\/em> dates approximately from 2.0 mya to 1 mya and is the only taxon from the genus to be discovered in South Africa. Several of these fossils are fragmentary in nature, distorted, and not well preserved because they have been recovered from quarry breccia using explosives. <em>P. robustus<\/em> features are neither as \u201chyper-robust\u201d as <em>P. boisei<\/em> nor as ancestral as <em>P. aethiopicus<\/em>; instead, they have been described as being less derived, more general features that are shared with both East African species (e.g., the sagittal crest and zygomatic flaring; Rak 1983; Walker and Leakey 1988). Enamel hypoplasia is also common in this species, possibly because of instability in the development of large, thick-enameled dentition.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 572px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-303 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.22.jpg\" alt=\"Four views of a beige-colored skull are shown on a black background.\" width=\"572\" height=\"619\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.22: SK 48, a Paranthropus robustus specimen, had less derived, more general features that were not as robust as P. boisei and not as ancestral as P. aethiopicus. Credit: a. <a class=\"rId208\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId209\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 anterior view<\/a> by \u00a9<a class=\"rId210\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId211\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId212\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId213\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId214\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 superior view<\/a> by \u00a9<a class=\"rId215\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId216\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId217\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a class=\"rId218\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId219\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 inferior view<\/a> by \u00a9<a class=\"rId220\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId221\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId222\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; d. <a class=\"rId223\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId224\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 lateral left view<\/a> by \u00a9<a class=\"rId225\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId226\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId227\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Comparisons between Gracile and Robust Australopiths<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Comparisons between gracile and robust australopithecines may indicate different phylogenetic groupings or parallel evolution in several species. In general, the robust australopithecines have large temporalis (chewing) muscles, as indicated by flaring zygomatic arches, sagittal crests, and robust mandibles (jawbones). Their hind dentition is large (megadont), with low cusps and thick enamel. Within the gracile australopithecines, researchers have debated the relatedness of the species, or even whether these species should be lumped together to represent more variable or polytypic species. Often researchers will attempt to draw chronospecific trajectories, with one taxon said to evolve into another over time.<\/span><\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Special Topic: The Taung Child<\/span><\/h2>\n<figure id=\"attachment_304\" aria-describedby=\"caption-attachment-304\" style=\"width: 570px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-304 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.23.jpg\" alt=\"An ancient skull in anterior and lateral views. One view shows an imprint of the brain.\" width=\"570\" height=\"285\" \/><figcaption id=\"caption-attachment-304\" class=\"wp-caption-text\">Figure 9.23: The Taung Child has a nearly complete face, mandible, and partial endocranial cast. Credit: a. <em>A<\/em><a class=\"rId230\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\"><em>ustralopithecus africanus<\/em><\/a><a class=\"rId231\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\">: Taung 1 anterior view<\/a> by \u00a9<a class=\"rId232\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId233\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId234\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId235\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\"><em>australopithecus africanus<\/em><\/a><a class=\"rId236\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\">: Taung 1 lateral right view<\/a> by \u00a9<a class=\"rId237\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId238\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId239\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">The well-known fossil of a juvenile <em>Australopithecine<\/em>, the \u201cTaung Child,\u201d was the first early hominin evidence ever discovered and was the first to demonstrate our common human heritage in Africa (Figure 9.23; Dart 1925). The tiny facial skeleton and natural endocast were discovered in 1924 by a local quarryman in the North West Province in South Africa and were painstakingly removed from the surrounding cement-like breccia by Raymond Dart using his wife\u2019s knitting needles. When first shared with the scientific community in 1925, it was discounted as being nothing more than a young monkey of some kind. Prevailing biases of the time made it too difficult to contemplate that this small-brained hominin could have anything to do with our own history. The fact that it was discovered in Africa simply served to strengthen this bias.<\/span><\/p>\n<\/div>\n<p>&nbsp;<\/p>\n<h2><span style=\"color: #000000\">Early Tool Use and Technology<br \/>\n<\/span><\/h2>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Early Stone Age Technology (ESA)<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1378\">Early Stone Age (ESA)<\/a><\/strong> marks the beginning of recognizable technology made by our human ancestors. Stone-tool (or <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1380\"><strong>lithic<\/strong><\/a>) technology is defined by the fracturing of rocks and the manufacture of tools through a process called <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1382\"> <strong>knapping<\/strong><\/a>. The Stone Age lasted for more than 3 million years and is broken up into chronological periods called the Early (ESA), Middle (MSA), and Later Stone Ages (LSA). Each period is further broken up into a different <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1384\">techno-complex<\/a><\/strong>, a term encompassing multiple <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1386\">assemblages<\/a><\/strong> (collections of artifacts) that share similar traits in terms of artifact production and morphology. The ESA spanned the largest technological time period of human innovation from over 3 million years ago to around 300,000 years ago and is associated almost entirely with hominin species prior to modern <em>Homo sapiens. <\/em>As the ESA advanced, stone tool makers (known as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1390\"><strong>knappers<\/strong><\/a>) began to change the ways they detached <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1388\">flakes<\/a><\/strong> and eventually were able to shape artifacts into functional tools. These advances in technology go together with the developments in human evolution and cognition, dispersal of populations across the African continent and the world, and climatic changes.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In order to understand the ESA, it is important to consider that not all assemblages are exactly the same within each techno-complex: one can have multiple phases and traditions at different sites (Lombard et al. 2012). However, there is an overarching commonality between them. Within stone tool assemblages, both flakes or <strong>cores<\/strong> (the rocks from which flakes are removed) are used as tools.<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1392\"> <strong>Large Cutting Tools (LCTs)<\/strong><\/a> are tools that are shaped to have functional edges. It is important to note that the information presented here is a small fraction of what is known about the ESA, and there are ongoing debates and discoveries within archaeology.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Currently, the oldest-known stone tools, which form the techno-complex the Lomekwian, date to 3.3 mya (Harmand et al. 2015; Toth 1985). They were found at a site called Lomekwi 3 in Kenya. This techno-complex is the most recently defined and pushed back the oldest-known date for lithic technology. There is only one known site thus far and, due to the age of the site, it is associated with species prior to <em>Homo<\/em>, such as <em>Kenyanthropus platyops.<\/em> Flakes were produced through indirect percussion, whereby the knappers held a rock and hit it against another rock resting on the ground. The pieces are very chunky and do not display the same fracture patterns seen in later techno-complexes. Lomekwian knappers likely aimed to get a sharp-edged piece on a flake, which would have been functional, although the specific function is currently unknown.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Stone tool use, however, is not only understood through the direct discovery of the tools. Cut marks on fossilized animal bones may illuminate the functionality of stone tools. In one controversial study in 2010, researchers argued that cut marks on a pair of animal bones from Dikika (Ethiopia), dated to 3.4 mya, were from stone tools. The discoverers suggested that they be more securely associated, temporally, with <em>Au. afarensis<\/em>. However, others have noted that these marks are consistent with teeth marks from crocodiles and other carnivores.<\/span><\/p>\n<figure style=\"width: 324px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image29-1.png\" alt=\"A technical line drawing of an Oldowan chopper.\" width=\"324\" height=\"275\" \/><figcaption class=\"wp-caption-text\">Figure 9.24: Some scholars believe that some genera explored in this chapter were capable of producing more complex stone tools (Oldowan). Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Olduwan_Industry_Chopper_2.jpeg\">Olduwan Industry Chopper 2<\/a> by Emmyanne29 is under a <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/legalcode\">CC0 1.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The Oldowan techno-complex is far more established in the scientific literature (Leakey 1971). It is called the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1394\">Oldowan<\/a><\/strong> because it was originally discovered in Olduvai Gorge, Tanzania, but the oldest assemblage is from Gona in Ethiopia, dated to 2.6 mya (Semaw 2000). The techno-complex is defined as a core and flake industry. Like the Lomekwian, there was an aim to get sharp-edged flakes, but this was achieved through a different production method. Knappers were able to actively hold or manipulate the core being knapped, which they could directly hit using a hammerstone. This technique is known as free-hand percussion, and it demonstrates an understanding of fracture mechanics. It has long been argued that the Oldowan hominins were skillful in tool manufacture.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Because Oldowan knapping requires skill, earlier researchers have attributed these tools to members of our genus, <em>Homo<\/em>. However, some have argued that these tools are in more direct association with hominins in the genera described in this chapter (Figure 9.24).<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Invisible Tool Manufacture and Use<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The vast majority of our understanding of these early hominins comes from fossils and reconstructed paleoenvironments. It is only from 3 mya when we can start \u201clooking into their minds\u201d and lifestyles by analyzing their manufacture and use of stone tools. However, the vast majority of tool use in primates (and, one can argue, in humans) is not with durable materials like stone. All of our extant great ape relatives have been observed using sticks, leaves, and other materials for some secondary purpose (to wade across rivers, to \u201cfish\u201d for termites, or to absorb water for drinking). It is possible that the majority of early hominin tool use and manufacture may be invisible to us because of this preservation bias.<\/span><\/p>\n<h2 class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000;background-color: #ff00ff\">Chapter Summary<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000;background-color: #ff00ff\">The fossil record of our earliest hominin relatives has allowed paleoanthropologists to unpack some of the mysteries of our evolution. We now know that traits associated with bipedalism evolved before other \u201chuman-like\u201d traits, even though the first hominins were still very capable of arboreal locomotion. We also know that, for much of this time, hominin taxa were diverse in the way they looked and what they ate, and they were widely distributed across the African continent. And we know that the environments in which these hominins lived underwent many changes over this time during several warming and cooling phases.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;background-color: #ff00ff\">Yet this knowledge has opened up many new mysteries. We still need to better differentiate some taxa. In addition, there are ongoing debates about why certain traits evolved and what they meant for the extinction of some of our relatives (like the robust australopiths). The capabilities of these early hominins with respect to tool use and manufacture is also still uncertain.<\/span><\/p>\n<h2 class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Hominin Species Summaries<br \/>\n<\/span><\/h2>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Sahelanthropus tchadensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">7 mya to 6 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Chad<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The initial discovery, made in 2001.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">360 cc average<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Smaller than in extant great apes; larger and pointier than in humans. Canines worn at the tips.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A short cranial base and a foramen magnum (hole in which the spinal cord enters the cranium) that is more humanlike in positioning; has been argued to indicate upright walking.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Currently little published postcranial material.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The extent to which this hominin was bipedal is currently heavily debated. If so, it would indicate an arboreal bipedal ancestor of hominins, not a knuckle-walker like chimpanzees.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Orrorin tugenensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">6 mya to 5.7 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Tugen Hills (Kenya)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Original discovery in 2000.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Smaller cheek teeth (molars and premolars) than even more recent hominins (i.e., derived), thick enamel, and reduced, but apelike, canines.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Not many found<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Fragmentary leg, arm, and finger bones have been found. Indicates bipedal locomotion.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Potential toolmaking capability based on hand morphology, but nothing found directly.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">This is the earliest species that clearly indicates adaptations for bipedal locomotion.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Ardipithecus kadabba<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">5.2 mya to 5.8 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Middle Awash (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Discovered by Yohannes Haile-Selassie in 1997.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Larger hind dentition than in modern chimpanzees. Thick enamel and larger canines than in later hominins.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A large hallux (big toe) bone indicates a bipedal \u201cpush off.\u201d<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Faunal evidence indicates a mixed grassland\/woodland environment.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Ardipithecus ramidus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">4.4 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Middle Awash region and Gona (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">A partial female skeleton nicknamed \u201cArdi\u201d (ARA-VP-6\/500) (found in 1994).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">300 cc to 350 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Little differences between the canines of males and females (small sexual dimorphism).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Midfacial projection, slightly prognathic. Cheekbones less flared and robust than in later hominins.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Ardi demonstrates a mosaic of ancestral and derived characteristics in the postcrania. For instance, an opposable big toe similar to chimpanzees (i.e., more ancestral), which could have aided in climbing trees effectively. However, the pelvis and hip show that she could walk upright (i.e., it is derived), supporting her hominin status.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">None directly associated<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">Over 110 specimens from Aramis<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus anamensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">4.2 mya to 3.8 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Turkana region (Kenya); Middle Awash (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A 2019 find from Ethiopia, named MRD.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">370 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Relatively large canines compared with more recent Australopithecines.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Projecting cheekbones and ancestral earholes.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lower limb bones (tibia and femur) indicate bipedality; arboreal features in upper limb bones (humerus) found.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Almost 100 specimens, representing over 20 individuals, have been found to date.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus afarensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">3.9 mya to 2.9 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Afar Region, Omo, Maka, Fejej, and Belohdelie (Ethiopia); Laetoli (Tanzania); Koobi Fora (Kenya)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lucy (discovery: 1974), Selam (Dikika Child, discovery: 2000), Laetoli Footprints (discovery: 1976).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">380 cc to 430 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Reduced canines and molars relative to great apes but larger than in modern humans.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Prognathic face, facial features indicate relatively strong chewing musculature (compared with <em>Homo<\/em>) but less extreme than in <em>Paranthropus<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clear evidence for bipedalism from lower limb postcranial bones. Laetoli Footprints indicate humanlike walking. Dikika Child bones indicate retained ancestral arboreal traits in the postcrania.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">None directly, but close in age and proximity to controversial cut marks at Dikika and early tools in Lomekwi.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Au. afarensis<\/em> is one of the oldest and most well-known australopithecine species and consists of a large number of fossil remains.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus bahrelghazali<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">3.6 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Chad<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">\u201cAbel,\u201d the holotype (discovery: 1995).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Arguably within range of variation of <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus prometheus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">3.7 mya (debated)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">Sterkfontein (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">\u201cLittle Foot\u201d (StW 573) (discovery: 1994)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">408 cc (Little Foot estimate)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">Heavy anterior dental wear patterns, relatively large anterior dentition and smaller hind dentition, similar to <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">Relatively larger brain size, robust zygomatic arch, and a flatter midface.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">The initial discovery of four ankle bones indicated bipedality.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">Highly debated new species designation.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus <\/em><em>deyiremada<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">3.5 mya to 3.3 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Woranso-Mille (Afar region, Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">First fossil mandible bones were discovered in 2011 in the Afar region of Ethiopia by Yohannes Haile-Selassie.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Smaller teeth with thicker enamel than seen in <em>Au. afarensis<\/em>, with a potentially hardier diet.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Larger mandible and more projecting cheekbones than in <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Contested species designation; arguably a member of <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Kenyanthopus<\/em><em> platyops<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">3.5 mya to 3.2 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Lake Turkana (Kenya)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">KNM\u2013WT 40000 (discovered 1999)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Difficult to determine but appears within the range of <em>Australopithecus afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Small molars\/dentition (<em>Homo<\/em>-like characteristic)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Flatter (i.e., orthognathic) face<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Some have associated the earliest tool finds from Lomekwi, Kenya, temporally (3.3 mya) and in close geographic proximity to this species\/specimen.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Taxonomic placing of this species is quite divided. The discoverers have argued that this species is ancestral to <em>Homo<\/em>, in particular to <em>Homo <\/em><em>ruldolfensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Australopithecus africanus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">3.3 mya to 2.1 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Sterkfontein, Taung, Makapansgat, Gladysvale (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Taung Child (discovery in 1994), \u201cMrs. Ples\u201d (discover in 1947), Little Foot (arguable; discovery in 1994).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">400 cc to 500 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Smaller teeth (derived) relative to <em>Au. afarensis<\/em>. Small canines with no diastema.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">A rounder skull compared with <em>Au. afarensis<\/em> in East Africa. A sloping face (ancestral).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Similar postcranial evidence for bipedal locomotion (derived pelvis) with retained arboreal locomotion, e.g., curved phalanges (fingers), as seen in <em>Au. afarensis.<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">None with direct evidence.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">A 2015 study noted that the trabecular bone morphology of the hand was consistent with forceful tool manufacture and use, suggesting potential early tool abilities.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus garhi<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.5 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Middle Awash (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">450 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Larger hind dentition than seen in other gracile Australopithecines.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A femur of a fragmentary partial skeleton, argued to belong to <em>Au. garhi<\/em>, indicates this species may be longer-limbed than <em>Au. afarensis<\/em>, although still able to move arboreally.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Crude stone tools resembling Oldowan (described later) have been found in association with <em>Au. garhi<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">This species is not well documented or understood and is based on only a few fossil specimens.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Paranthropus aethiopicus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.7 mya to 2.3 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">West Turkana (Kenya); Laetoli (Tanzania); Omo River Basin (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The \u201cBlack Skull\u201d (KNM\u2013WT 17000) (discovery 1985).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain Size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">410 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>P. aethiopicus<\/em> has the shared derived traits of large flat premolars and molars, although few teeth have been found.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Large flaring zygomatic arches for accommodating large chewing muscles (the temporalis muscle), a sagittal crest for increased muscle attachment of the chewing muscles to the skull, and a robust mandible and supraorbital torus (brow ridge).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A proximal tibia indicates bipedality and similar size to <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The \u201cBlack Skull\u201d is so called because of the mineral manganese that stained it black during fossilization.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Paranthropus boisei<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.4 mya to 1.4 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Koobi Fora, West Turkana, and Chesowanja (Kenya); Malema-Chiwondo (Malawi), Olduvai Gorge and Peninj (Tanzania); and Omo River basin and Konso (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">\u201cZinj,\u201d or sometimes \u201cNutcracker Man\u201d (OH5), in 1959 by Mary Leakey. The Peninj mandible from Tanzania, found in 1964 by Kimoya Kimeu.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">500 cc to 550 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Very large, flat posterior dentition (largest of all hominins currently known). Much smaller anterior dentition. Very thick dental enamel.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Indications of very large chewing muscles (e.g., flaring zygomatic arches and a large sagittal crest).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Evidence for high variability and sexual dimorphism, with estimates of males at 1.37 meters tall and females at 1.24 meters.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Richard Leakey and Bernard Wood have both suggested that<em> P. boisei<\/em> could have made and used stone tools. Tools dated to 2.5 mya in Ethiopia have been argued to possibly belong to this species.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Despite the cranial features of <em>P. boisei<\/em> indicating a tough diet of tubers, nuts, and seeds, isotopes indicate a diet high in C4 foods (e.g., grasses, such as sedges). This differs from what is seen in<em> P. robustus<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Australopithecus sediba<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">1.97 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Malapa Fossil Site (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Karabo (MH1) (discovery in 2008)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">420 cc to 450 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Small dentition with Australopithecine cusp-spacing.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Small brain size (<em>Australopithecus<\/em>-like) but gracile mandible (<em>Homo<\/em>-like).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Scientists have interpreted this mixture of traits (such as a robust ankle but evidence for an arch in the foot) as a transitional phase between a body previously adapted to arborealism (tree climbing, particularly in evidence from the bones of the wrist) to one that adapted to bipedal ground walking.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">None of direct association, but some have argued that a modern hand morphology (shorter fingers and a longer thumb) means that adaptations to tool manufacture and use may be present in this species.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">It was first discovered through a clavicle bone in 2008 by nine-year-old Matthew Berger, son of paleoanthropologist Lee Berger.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Paranthropus robustus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.3 mya to 1 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kromdraai B, Swartkrans, Gondolin, Drimolen, and Coopers Cave (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">SK48 (original skull)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">410 cc to 530 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Large posterior teeth with thick enamel, consistent with other Robust Australopithecines. Enamel hypoplasia is also common in this species, possibly because of instability in the development of large, thick enameled dentition.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>P. robustus<\/em> features are neither as \u201chyper-robust\u201d as <em>P. boisei<\/em> or as ancestral in features as <em>P. aethiopicus<\/em>. They have been described as less derived, more general features that are shared with both East African species (e.g., the sagittal crest and zygomatic flaring).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Reconstructions indicate sexual dimorphism.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Several of these fossils are fragmentary in nature, distorted, and not well preserved, because they have been recovered from quarry breccia using explosives.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\"><strong><span style=\"color: #000000\">Review Questions<br \/>\n<\/span><\/strong><\/h2>\n<ul>\n<li class=\"import-Normal\"><span style=\"color: #000000\">What is the difference between a \u201cderived\u201d versus an \u201cancestral\u201d trait? Give an example of both, seen in <em>Au. afarensis<\/em>.<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">Which of the paleoenvironment hypotheses have been used to describe early hominin diversity, and which have been used to describe bipedalism?<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">Which anatomical features for bipedalism do we see in early hominins?<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">Describe the dentition of gracile and robust australopithecines. What might these tell us about their diets?<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">List the hominin species argued to be associated with stone tool technologies. Are you convinced of these associations? Why\/why not?<\/span><\/li>\n<\/ul>\n<\/div>\n<h2><span style=\"color: #000000\">Key Terms<\/span><\/h2>\n<p><span style=\"color: #000000\"><strong>Arboreal:<\/strong> Related to trees or woodland.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Aridification:<\/strong> Becoming increasingly arid or dry, as related to the climate or environment.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Aridity Hypothesis:<\/strong> The hypothesis that long-term aridification and expansion of savannah biomes were drivers in diversification in early hominin evolution.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Assemblage:<\/strong> A collection demonstrating a pattern. Often pertaining to a site or region.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Bipedalism:<\/strong> The locomotor ability to walk on two legs.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Breccia:<\/strong> Hard, calcareous sedimentary rock.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Canines:<\/strong> The pointy teeth just next to the incisors, in the front of the mouth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cheek teeth:<\/strong> Or hind dentition (molars and premolars).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Chronospecies:<\/strong> Species that are said to evolve into another species, in a linear fashion, over time.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Clade:<\/strong> A group of species or taxa with a shared common ancestor.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cladistics:<\/strong> The field of grouping organisms into those with shared ancestry.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Context:<\/strong> As pertaining to palaeoanthropology, this term refers to the place where an artifact or fossil is found.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cores:<\/strong> The remains of a rock that has been flaked or knapped.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cusps:<\/strong> The ridges or \u201cbumps\u201d on the teeth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Dental formula:<\/strong> A technique to describe the number of incisors, canines, premolars, and molars in each quadrant of the mouth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Derived traits:<\/strong> Newly evolved traits that differ from those seen in the ancestor.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Diastema:<\/strong> A tooth gap between the incisors and canines.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Early Stone Age (ESA):<\/strong> The earliest-described archaeological period in which we start seeing stone-tool technology.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>East African Rift System (EARS):<\/strong> This term is often used to refer to the Rift Valley, expanding from Malawi to Ethiopia. This active geological structure is responsible for much of the visibility of the paleoanthropological record in East Africa.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Enamel:<\/strong> The highly mineralized outer layer of the tooth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Encephalization:<\/strong> Expansion of the brain.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Extant:<\/strong> Currently living\u2014i.e., not extinct.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Fallback foods:<\/strong> Foods that may not be preferred by an animal (e.g., foods that are not nutritionally dense) but that are essential for survival in times of stress or scarcity.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Fauna:<\/strong> The animals of a particular region, habitat, or geological period.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Faunal assemblages:<\/strong> Collections of fossils of the animals found at a site.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Faunal turnover:<\/strong> The rate at which species go extinct and are replaced with new species.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Flake:<\/strong> The piece knocked off of a stone core during the manufacture of a tool, which may be used as a stone tool.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Flora:<\/strong> The plants of a particular region, habitat, or geological period.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Folivorous:<\/strong> Foliage-eating.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Foramen magnum:<\/strong> The large hole (foramen) at the base of the cranium, through which the spinal cord enters the skull.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Fossil:<\/strong> The remains or impression of an organism from the past.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Frugivorous:<\/strong> Fruit-eating.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Generalist:<\/strong> A species that can thrive in a wide variety of habitats and can have a varied diet.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Glacial:<\/strong> Colder, drier periods during an ice age when there is more ice trapped at the poles.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Gracile:<\/strong> Slender, less rugged, or pronounced features.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hallux:<\/strong> The big toe.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Holotype:<\/strong> A single specimen from which a species or taxon is described or named.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hominin:<\/strong> A primate category that includes humans and our fossil relatives since our divergence from extant great apes.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Honing P3:<\/strong> The mandibular premolar alongside the canine (in primates, the P3), which is angled to give space for (and sharpen) the upper canines.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hyper-robust:<\/strong> Even more robust than considered normal in the Paranthropus genus.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hypodigm:<\/strong> A sample (here, fossil) from which researchers extrapolate features of a population.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Incisiform:<\/strong> An adjective referring to a canine that appears more incisor-like in morphology.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Incisors:<\/strong> The teeth in the front of the mouth, used to bite off food.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Interglacial:<\/strong> A period of milder climate in between two glacial periods.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Isotopes:<\/strong> Two or more forms of the same element that contain equal numbers of protons but different numbers of neutrons, giving them the same chemical properties but different atomic masses.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Knappers:<\/strong> The people who fractured rocks in order to manufacture tools.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Knapping:<\/strong> The fracturing of rocks for the manufacture of tools.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Large Cutting Tool (LCT):<\/strong> A tool that is shaped to have functional edges.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Last Common Ancestor (LCA):<\/strong> The hypothetical final ancestor (or ancestral population) of two or more taxa before their divergence.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Lithic:<\/strong> Relating to stone (here to stone tools).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Lumbar lordosis:<\/strong> The inward curving of the lower (lumbar) parts of the spine. The lower curve in the human S-shaped spine.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Lumpers:<\/strong> Researchers who prefer to lump variable specimens into a single species or taxon and who feel high levels of variation is biologically real.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Megadont:<\/strong> An organism with extremely large dentition compared with body size.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Metacarpals:<\/strong> The long bones of the hand that connect to the phalanges (finger bones).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Molars:<\/strong> The largest, most posterior of the hind dentition.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Monophyletic:<\/strong> A taxon or group of taxa descended from a common ancestor that is not shared with another taxon or group.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Morphology:<\/strong> The study of the form or size and shape of things; in this case, skeletal parts.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Mosaic evolution:<\/strong> The concept that evolutionary change does not occur homogeneously throughout the body in organisms.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Obligate bipedalism:<\/strong> Where the primary form of locomotion for an organism is bipedal.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Occlude:<\/strong> When the teeth from the maxilla come into contact with the teeth in the mandible.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Oldowan:<\/strong> Lower Paleolithic, the earliest stone tool culture.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Orthognathic:<\/strong> The face below the eyes is relatively flat and does not jut out anteriorly.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Paleoanthropologists:<\/strong> Researchers that study human evolution.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Paleoenvironment:<\/strong> An environment from a period in the Earth\u2019s geological past.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Parabolic:<\/strong> Like a parabola (parabola-shaped).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Phalanges:<\/strong> Long bones in the hand and fingers.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Phylogenetics:<\/strong> The study of phylogeny.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Phylogeny:<\/strong> The study of the evolutionary relationships between groups of organisms.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Pliocene:<\/strong> A geological epoch between the Miocene and Pleistocene.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Polytypic:<\/strong> In reference to taxonomy, having two or more group variants capable of interacting and breeding biologically but having morphological population differences.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Postcranium:<\/strong> The skeleton below the cranium (head).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Premolars:<\/strong> The smallest of the hind teeth, behind the canines.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Procumbent:<\/strong> In reference to incisors, tilting forward.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Prognathic:<\/strong> In reference to the face, the area below the eyes juts anteriorly.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Quaternary Ice Age:<\/strong> The most recent geological time period, which includes the Pleistocene and Holocene Epochs and which is defined by the cyclicity of increasing and decreasing ice sheets at the poles.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Relative dating:<\/strong> Dating techniques that refer to a temporal sequence (i.e., older or younger than others in the reference) and do not estimate actual or absolute dates.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Robust:<\/strong> Rugged or exaggerated features.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Site:<\/strong> A place in which evidence of past societies\/species\/activities may be observed through archaeological or paleontological practice.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Specialist:<\/strong> A specialist species can thrive only in a narrow range of environmental conditions or has a limited diet.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Splitters:<\/strong> Researchers who prefer to split a highly variable taxon into multiple groups or species.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Taxa:<\/strong> Plural of taxon, a taxonomic group such as species, genus, or family.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Taxonomy:<\/strong> The science of grouping and classifying organisms.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Techno-complex:<\/strong> A term encompassing multiple assemblages that share similar traits in terms of artifact production and morphology.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Thermoregulation:<\/strong> Maintaining body temperature through physiologically cooling or warming the body.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Ungulates:<\/strong> Hoofed mammals\u2014e.g., cows and kudu.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Volcanic tufts:<\/strong> Rock made from ash from volcanic eruptions in the past.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Valgus knee:<\/strong> The angle of the knee between the femur and tibia, which allows for weight distribution to be angled closer to the point above the center of gravity (i.e., between the feet) in bipeds.<\/span><\/p>\n<h2 class=\"import-Normal\"><strong><span style=\"color: #000000\">About the Authors<br \/>\n<\/span><\/strong><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image31.jpg\" alt=\"A woman with short blonde hair smiles at the camera.\" width=\"311\" height=\"311\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Kerryn Warren, Ph.D.<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Grad Coach International, <a class=\"rId245\" style=\"color: #000000\" href=\"mailto:kerryn.warren@gmail.com\">kerryn.warren@gmail.com<\/a><\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kerryn Warren is a dissertation coach at Grad Coach International and is passionate about stimulating research thinking in students of all levels. She has lectured on multiple topics, including archaeology and human evolution, with her research and science communication interests including hybridization in the hominin fossil record (stemming from research from her Ph.D.) and understanding how evolution is taught in South African schools. She also worked as one of the \u201cUnderground Astronauts,\u201d selected to excavate <em>Homo naledi <\/em>remains from the Rising Star Cave System in the Cradle of Humankind.<\/span><\/p>\n<p>&nbsp;<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image11-2.jpg\" alt=\"A woman with short brown hair smiles at the camera.\" width=\"312\" height=\"306\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">K. Lindsay Hunter, M.A., Ph.D. candidate<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">CARTA, k.lindsay.hunter@gmail.com<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lindsay Hunter is a trained palaeoanthropologist who uses her more than 15 years of experience to make sense of the distant past of our species to build a better future. She received her master\u2019s degree in biological anthropology from the University of Iowa and is completing her Ph.D. in archaeology at the University of the Witwatersrand in Johannesburg, South Africa. She has studied fossil and human bone collections across five continents with major grant support from the National Science Foundation (United States) and the Wenner-Gren Foundation for Anthropological Research. As a National Geographic Explorer, Lindsay developed and managed the National Geographic\u2013sponsored Umsuka Public Palaeoanthropology Project in the Cradle of Humankind World Heritage Site (CoH WHS) in South Africa from within Westbury Township, Johannesburg, between 2016\u20132019. She currently serves as the Community Engagement &amp; Advancement Director for CARTA: The UC San Diego\/Salk Institute Center for Academic Research and Training in Anthropogeny in La Jolla, California.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image25-2.jpg\" alt=\"A woman with black hair stands in a hole in the ground.\" width=\"295\" height=\"339\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Navashni Naidoo, M.Sc.<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">University of Cape Town, nnaidoo2@illinois.edu<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Navashni Naidoo is a researcher at Nelson Mandela University, lecturing on physical geology. She completed her Master\u2019s in Science in Archaeology in 2017 at the University of Cape Town. Her research interests include developing paleoenvironmental proxies suited to the African continent, behavioral ecology, and engaging with community-driven archaeological projects. She has excavated at Stone Age sites across Southern Africa and East Africa. Navashni is currently pursuing a PhD in the Department of Anthropology at the University of Illinois.<\/span><\/p>\n<p>&nbsp;<\/p>\n<p>&nbsp;<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image47.jpg\" alt=\"A man with black hair and dark brown eyes looks at the camera. \" width=\"294\" height=\"294\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Silindokuhle Mavuso, M.Sc.<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">University of Witwatersrand, S.muvaso@ru.ac.za<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Silindokuhle has always been curious about the world around him and how it has been shaped. He is a lecturer at Rhodes University of Witwatersrand (Wits), and conducts research on palaeoenvironmental reconstruction and change of the northeastern Turkana Basin\u2019s Pleistocene sequence. Silindokuhle began his education with a B.Sc. (Geology, Archaeology, and Environmental and Geographical Sciences) from the University of Cape Town before moving to Wits for a B.Sc. Honors (geology and paleontology) and M.Sc. in geology. He is currently concluding his PhD Studies. During this time, he has gained more training as a Koobi Fora Fieldschool fellow (Kenya) as well as an Erasmus Mundus scholar (France). Silindokuhle is a Plio-Pleistocene geologist with a specific focus on identifying and explaining past environments that are associated with early human life and development through time. He is interested in a wide range of disciplines such as micromorphology, sedimentology, geochemistry, geochronology, and sequence stratigraphy. He has worked with teams from significant eastern and southern African hominid sites including Elandsfontein, Rising Star, Sterkfontein, Gondolin, Laetoli, Olduvai, and Koobi Fora.<\/span><\/p>\n<h2 class=\"import-Normal\"><strong><span style=\"color: #000000\">For Further Exploration<br \/>\n<\/span><\/strong><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/humanorigins.si.edu\/evidence\">The Smithsonian Institution website<\/a> hosts descriptions of fossil species, an interactive timeline, and much more.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.maropeng.co.za\/content\/page\/human-evolution\">The Maropeng Museum website<\/a> hosts a wealth of information regarding South African Fossil Bearing sites in the Cradle of Humankind<strong>.<\/strong><\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/perot-museum.imgix.net\/2019-08-naledi-sediba-quick-comparison.pdf\">This quick comparison between <em>Homo naledi<\/em> and <em>Australopithecus sediba<\/em><\/a> from the Perot Museum.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.dropbox.com\/s\/l1d2hv42psj21y9\/Braided%20Stream-1920.mp4?dl=0\">This explanation of the braided stream<\/a> by the Perot Museum.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.hetmp.com\/\">A collation of 3-D files for visualizing<\/a> (or even 3-D printing) for homes, schools, and universities.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.pbslearningmedia.org\/resource\/tdc02.sci.life.evo.lp_humanevo\/human-evolution.\">PBS learning materials<\/a>, including videos and diagrams of the Laetoli footprints, bipedalism, and fossils.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A wealth of <a href=\"https:\/\/australianmuseum.net.au\/learn\/science\/human-evolution\/\">information from the Australian Museum website<\/a>, including species descriptions, family trees, and explanations of bipedalism and diet<strong>.<\/strong><\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\"><strong>References<\/strong><\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Alemseged, Zeresenay, Fred Spoor, William H. Kimbel, Ren\u00e9 Bobe, Denis Geraads, Denn\u00e9 Reed, and Jonathan G. Wynn. 2006. \u201cA Juvenile Early Hominin Skeleton from Dikika, Ethiopia.\u201d <em>Nature<\/em> 443 (7109): 296\u2013301.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Asfaw, Berhane, Tim White, Owen Lovejoy, Bruce Latimer, Scott Simpson, and Gen Suwa. 1999. \u201c<em>Australopithecus garhi<\/em>: A New Species of Early Hominid from Ethiopia.\u201d <em>Science<\/em> 284 (5414): 629\u2013635.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Behrensmeyer, Anna K., Nancy E. Todd, Richard Potts, and Geraldine E. McBrinn. 1997. \u201cLate Pliocene Faunal Turnover in the Turkana Basin, Kenya, and Ethiopia.\u201d <em>Science<\/em> 278 (5343): 637\u2013640.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Berger, Lee R., Darryl J. De Ruiter, Steven E. Churchill, Peter Schmid, Kristian J. Carlson, Paul HGM Dirks, and Job M. Kibii. 2010. \u201c<em>Australopithecus sediba<\/em>: A New Species of <em>Homo<\/em>-like Australopith from South Africa.\u201d <em>Science<\/em> 328 (5975): 195\u2013204.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Bobe, Ren\u00e9, and Anna K. Behrensmeyer. 2004. \u201cThe Expansion of Grassland Ecosystems in Africa in Relation to Mammalian Evolution and the Origin of the Genus <em>Homo<\/em>.\u201d <em>Palaeogeography, Palaeoclimatology, Palaeoecology<\/em> 207 (3\u20134): 399\u2013420.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Brain, C. K. 1967. \u201cThe Transvaal Museum's Fossil Project at Swartkrans.\u201d <em>South African Journal of Science<\/em> 63 (9): 378\u2013384.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1938a. \u201cMore Discoveries of Australopithecus.\u201d <em>Nature<\/em> 141 (1): 828\u2013829.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1938b. \u201cThe Pleistocene Anthropoid Apes of South Africa.\u201d <em>Nature<\/em> 142 (3591): 377\u2013379.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1947. \u201cDiscovery of a New Skull of the South African Ape-Man, Plesianthropus.\u201d <em>Nature<\/em> 159 (4046): 672.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1950. \u201cThe Genera and Species of the South African Fossil Ape-Man.\u201d <em>American Journal of Physical Anthropology<\/em> 8 (1): 1\u201314.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Brunet, Michel, Alain Beauvilain, Yves Coppens, Emile Heintz, Aladji HE Moutaye, and David Pilbeam. 1995. \u201cThe First Australopithecine 2,500 Kilometers West of the Rift Valley (Chad).\u201d <em>Nature<\/em> 378 (6554): 275\u2013273.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Cerling, Thure E., Jonathan G. Wynn, Samuel A. Andanje, Michael I. Bird, David Kimutai Korir, Naomi E. Levin, William Mace, Anthony N. Macharia, Jay Quade, and Christopher H. Remien. 2011. \u201cWoody Cover and Hominin Environments in the Past 6 Million Years.\u201d <em>Nature<\/em> 476, no. 7358 (2011): 51-56..<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clarke, Ronald J. 1998. \u201cFirst Ever Discovery of a Well-Preserved Skull and Associated Skeleton of <em>Australopithecus<\/em>.\u201d <em>South African Journal of Science<\/em> 94 (10): 460\u2013463.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Clarke, Ronald J. 2013. \u201cAustralopithecus from Sterkfontein Caves, South Africa.\u201d In <em>The Paleobiology of Australopithecus<\/em>, edited by K. E. Reed, J. G. Fleagle, and R. E. Leakey, 105\u2013123. Netherlands: Springer.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clarke, Ronald J., and Kathleen Kuman. 2019. \u201cThe Skull of StW 573, a 3.67 Ma Australopithecus Prometheus Skeleton from Sterkfontein Caves, South Africa.\u201d <em>Journal of <\/em><em>H<\/em><em>uman <\/em><em>E<\/em><em>volution<\/em> 134: 102634.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clarke, R. J., and P. V. Tobias. 1995. \u201cSterkfontein Member 2 Foot Bones of the Oldest South African Hominid.\u201d <em>Science<\/em> 269 (5223): 521\u2013524.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Constantino, P. J., and B. A. Wood. 2004. \u201cParanthropus Paleobiology\u201d. In <em>Miscelanea en <\/em><em>Homenae<\/em><em> a Emiliano Aguirre<\/em><em>,<\/em> <em>v<\/em><em>olumen III: Paleoantropologia<\/em>, edited by E. G. P\u00e9rez and S. R. Jara, 136\u2013151. Alcal\u00e1 de Henares: Museo Arqueologico Regional.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Constantino, P. J., and B. A. Wood. 2007. \u201cThe Evolution of Zinjanthropus boisei.\u201d <em>Evolutionary Anthropology: <\/em><em>Issues, News, and Reviews<\/em> 16 (2): 49\u201362.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Dart, Raymond A. 1925. \u201cAustralopithecus africanus, the Man-Ape of South Africa.\u201d <em>Nature<\/em> 115: 195\u2013199.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Darwin, Charles. 1871. <em>The Descent of Man: And Selection in Relation to Sex<\/em>. London: J. Murray.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Daver, Guillaume, F. Guy, Hassane Ta\u00efsso Mackaye, Andossa Likius, J-R. Boisserie, Abderamane Moussa, Laurent Pallas, Patrick Vignaud, and N\u00e9koulnang D. Clarisse. 2022. \"Postcranial Evidence of Late Miocene Hominin Bipedalism in Chad.\" <em>Nature<\/em> 609 (7925): 94\u2013100.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Heinzelin, Jean de, J. Desmond Clark, Tim White, William Hart, Paul Renne, Giday WoldeGabriel, Yonas Beyene, and Elisabeth Vrba. 1999. \u201cEnvironment and Behavior of 2.5-Million-Year-Old Bouri Hominids.\u201d <em>Science<\/em> 284 (5414): 625\u2013629.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">DeMenocal, Peter B. D. 2004. \u201cAfrican Climate Change and Faunal Evolution during the Pliocene\u2013Pleistocene.\u201d <em>Earth and Planetary Science Letters<\/em> 220 (1\u20132): 3\u201324.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">DeMenocal, Peter B. D. and J. Bloemendal, J. 1995. \u201cPlio-Pleistocene Climatic Variability in Subtropical Africa and the Paleoenvironment of Hominid Evolution: A Combined Data-Model Approach.\u201d In <em>Paleoclimate and Evolution, with Emphasis on Human Origins<\/em>, edited by E. S. Vrba, G. H. Denton, T. C. Partridge, and L. H. Burckle, 262\u2013288. New Haven: Yale University Press.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Dirks, Paul HGM, Job M. Kibii, Brian F. Kuhn, Christine Steininger, Steven E. Churchill, Jan D. Kramers, Robyn Pickering, Daniel L. Farber, Anne-Sophie M\u00e9riaux, Andy I. R. Herries, Geoffrey C. P. King, And Lee R. Berger. 2010. \u201cGeological Setting and Age of <em>Australopithecus sediba<\/em> from Southern Africa.\u201d <em>Science<\/em> 328 (5975): 205\u2013208.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Faith, J. Tyler, and Anna K. Behrensmeyer. 2013. \u201cClimate Change and Faunal Turnover: Testing the Mechanics of the Turnover-Pulse Hypothesis with South African Fossil Data.\u201d <em>Paleobiology<\/em> 39 (4): 609\u2013627.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Grine, Frederick E. 1988. \u201cNew Craniodental Fossils of <em>Paranthropus<\/em> from the Swartkrans Formation and Their Significance in \u2018Robust\u2019 Australopithecine Evolution.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by F. E. Grine, 223\u2013243. New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Grine, Frederick E., Carrie S. Mongle, John G. Fleagle, and Ashley S. Hammond. 2022. \"The Taxonomic Attribution of African Hominin Postcrania from the Miocene through the Pleistocene: Associations and Assumptions.\" <em>Journal of Human Evolution<\/em> 173: 103255.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Haile-Selassie, Yohannes, Luis Gibert, Stephanie M. Melillo, Timothy M. Ryan, Mulugeta Alene, Alan Deino, Naomi E. Levin, Gary Scott, and Beverly Z. Saylor. 2015. \u201cNew Species from Ethiopia Further Expands Middle Pliocene Hominin Diversity.\u201d <em>Nature<\/em> 521 (7553): 432\u2013433.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Haile-Selassie, Yohannes, Stephanie M. Melillo, Antonino Vazzana, Stefano Benazzi, and Timothy M. Ryan. 2019. \u201cA 3.8-Million-Year-Old Hominin Cranium from Woranso-Mille, Ethiopia.\u201d <em>Nature<\/em> 573 (7773): 214-219.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Harmand, Sonia, Jason E. Lewis, Craig S. Feibel, Christopher J. Lepre, Sandrine Prat, Arnaud Lenoble, Xavier Bo\u00ebs et al. 2015. \u201c3.3-Million-Year-Old Stone Tools from Lomekwi3, West Turkana, Kenya.\u201d <em>Nature<\/em> 521 (7552): 310\u2013316.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Hay, Richard L. 1990. \u201cOlduvai Gorge: A Case History in the Interpretation of Hominid Paleoenvironments.\u201d In <em>East Africa: Establishment of a Geologic Framework for Paleoanthropology<\/em>, edited by L. Laporte, 23\u201337<em>.<\/em> Boulder: Geological Society of America.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Hay, Richard L., and Mary D. Leakey. 1982. \u201cThe Fossil Footprints of Laetoli.\u201d <em>Scientific American<\/em> 246 (2): 50\u201357.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Hlazo, Nomawethu. 2015. \u201cParanthropus: Variation in Cranial Morphology.\u201d Honours thesis, Archaeology Department, University of Cape Town, Cape Town.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Hlazo, Nomawethu. 2018. \u201cVariation and the Evolutionary Drivers of Diversity in the Genus <em>Paranthropus<\/em>.\u201d Master\u2019s thesis, Archaeology Department, University of Cape Town, Cape Town.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Johanson, D. C., T. D. White, and Y. Coppens. 1978. \u201cA New Species of the Genus <em>Australopithecus<\/em> (Primates: Hominidae) from the Pliocene of East Africa.\u201d <em>Kirtlandia<\/em> 28: 1\u201314.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kimbel, William H. 2015. \u201cThe Species and Diversity of Australopiths.\u201d In <em>Handbook of Paleoanthropology<\/em>, 2nd ed., edited by T. Hardt, 2071\u20132105. Berlin: Springer.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kimbel, William H., and Lucas K. Delezene. 2009. \u201c\u2018Lucy\u2019 Redux: A Review of Research on <em>Australopithecus afarensis<\/em>.\u201d <em>American <\/em><em>J<\/em><em>ournal of <\/em><em>P<\/em><em>hysical <\/em><em>A<\/em><em>nthropology<\/em> 140 (S49): 2\u201348.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Kingston, John D. 2007. \u201cShifting Adaptive Landscapes: Progress and Challenges in Reconstructing Early Hominid Environments.\u201d <em>American Journal of Physical Anthropology<\/em> 134 (S45): 20\u201358.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Kingston, John D., and Terry Harrison. 2007. \u201cIsotopic Dietary Reconstructions of Pliocene Herbivores at Laetoli: Implications for Early Hominin Paleoecology.\u201d <em>Palaeogeography, Palaeoclimatology, Palaeoecology<\/em> 243 (3\u20134): 272\u2013306.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Leakey, Louis S. B. 1959. \u201cA New Fossil Skull from Olduvai.\u201d <em>Nature<\/em> 184 (4685): 491\u2013493.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Leakey, Mary 1971. <em>Olduvai Gorge<\/em>, Vol. 3. Cambridge: Cambridge University Press.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Leakey, Mary D., and Richard L. Hay. 1979. \u201cPliocene Footprints in the Laetoli Beds at Laetoli, Northern Tanzania.\u201d <em>Nature<\/em> 278 (5702): 317\u2013323.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Leakey, Meave G., Craig S. Feibel, Ian McDougall, and Alan Walker. 1995. \u201cNew Four\u2013Million-Year-Old Hominid Species from Kanapoi and Allia Bay, Kenya.\u201d <em>Nature<\/em> 376 (6541): 565\u2013571.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Meave G., Fred Spoor, Frank H. Brown, Patrick N. Gathogo, Christopher Kiarie, Louise N. Leakey, and Ian McDougall. 2001. \u201cNew Hominin Genus from Eastern Africa Shows Diverse Middle Pliocene Lineages.\u201d <em>Nature<\/em> 410 (6827): 433\u2013440.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lebatard, Anne-Elisabeth, Didier L. Bourl\u00e8s, Philippe Duringer, Marc Jolivet, R\u00e9gis Braucher, Julien Carcaillet, Mathieu Schuster et al. 2008. \u201cCosmogenic Nuclide Dating of <em>Sahelanthropus tchadensis<\/em> and <em>Australopithecus bahrelghazali<\/em>: Mio-Pliocene Hominids from Chad.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 105 (9): 3226\u20133231.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lee-Thorp, Julia. 2011. \u201cThe Demise of \u2018Nutcracker Man.\u2019\u201d <em>Proceedings of the National Academy of Sciences<\/em> 108 (23): 9319\u20139320.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lombard, Marlize, L. Y. N. Wadley, Janette Deacon, Sarah Wurz, Isabelle Parsons, Moleboheng Mohapi, Joane Swart, and Peter Mitchell. 2012. \u201cSouth African and Lesotho Stone Age Sequence Updated.\u201d <em>The South African Archaeological Bulletin<\/em> 67 (195): 123\u2013144.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Maslin, Mark A., Chris M. Brierley, Alice M. Milner, Susanne Shultz, Martin H. Trauth, and Katy E. Wilson. 2014. \u201cEast African Climate Pulses and Early Human Evolution.\u201d <em>Quaternary Science Reviews<\/em> 101: 1\u201317.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">McHenry, Henry M. 2009. \u201cHuman Evolution.\u201d In <em>Evolution: The First Four Billion Years<\/em>, edited by M. Ruse and J. Travis, 256\u2013280. Cambridge: The Belknap Press of Harvard University Press..<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Patterson, Bryan, and William W. Howells. 1967. \u201cHominid Humeral Fragment from Early Pleistocene of Northwestern Kenya.\u201d <em>Science<\/em> 156 (3771): 64\u201366.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Pickering, Robyn, and Jan D. Kramers. 2010. \u201cRe-appraisal of the Stratigraphy and Determination of New U-Pb Dates for the Sterkfontein Hominin Site.\u201d <em>Journal of Human Evoluti<\/em><em>on<\/em> 59 (1): 70\u201386.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Potts, Richard. 1998. \u201cEnvironmental Hypotheses of Hominin Evolution.\u201d <em>American Journal of Physical Anthropology<\/em> 107 (S27): 93\u2013136.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Potts, Richard. 2013. \u201cHominin Evolution in Settings of Strong Environmental Variability.\u201d <em>Quaternary Science Reviews<\/em> 73: 1\u201313.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Rak, Yoel. 1983. <em>The <\/em><em>A<\/em><em>ustralopithecine <\/em><em>F<\/em><em>ace<\/em>. New York: Academic Press.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Rak, Yoel. 1988. \u201cOn Variation in the Masticatory System of <em>Australopithecus boisei<\/em>.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by M. Ruse and J. Travis, 193\u2013198<em>.<\/em> New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Semaw, Sileshi. 2000. \u201cThe World\u2019s Oldest Stone Artefacts from Gona, Ethiopia: Their Implications for Understanding Stone Technology and Patterns of Human Evolution between 2.6 Million Years Ago and 1.5 Million Years Ago.\u201d <em>Journal of Archaeological Science<\/em> 27(12): 1197\u20131214.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Shipman, Pat. 2002. <em>The Man Who Found the Missing Link: Eug<\/em><em>e<\/em><em>ne Dubois and <\/em><em>h<\/em><em>is Lifelong Quest to Prove Darwin Right<\/em>. New York: Simon &amp; Schuster.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Spoor, Fred. 2015. \u201cPalaeoanthropology: The Middle Pliocene Gets Crowded.\u201d<em> Nature<\/em> 521 (7553): 432\u2013433.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Strait, David S., Frederick E. Grine, and Marc A. Moniz. 1997. A Reappraisal of Early Hominid Phylogeny.\u201d <em>Journal of <\/em><em>H<\/em><em>uman <\/em><em>E<\/em><em>volution<\/em> 32 (1): 17\u201382.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Thackeray, J. Francis. 2000. \u201c\u2018Mrs. Ples\u2019 from Sterkfontein: Small Male or Large Female?\u201d <em>The South African Archaeological <\/em><em>Bulletin<\/em> 55: 155\u2013158.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Thackeray, J. Francis, Jos\u00e9 Braga, Jacques Treil, N. Niksch, and J. H. Labuschagne. 2002. \u201c\u2018Mrs. Ples\u2019 (Sts 5) from Sterkfontein: An Adolescent Male?\u201d <em>South African Journal of Science<\/em> 98 (1\u20132): 21\u201322.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Toth, Nicholas. 1985. \u201cThe Oldowan Reassessed.\u201d <em>Journal of Archaeological Science<\/em>\u00a012 (2): 101\u2013120.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Vrba, E. S. 1988. \u201cLate Pliocene Climatic Events and Hominid Evolution.\u201d In <em>The <\/em><em>E<\/em><em>volutionary <\/em><em>H<\/em><em>istory of the <\/em><em>R<\/em><em>obust Australopithecines<\/em>, edited by F. E. Grine, 405\u2013426. New York: Aldine.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Vrba, Elisabeth S. 1998. \u201cMultiphasic Growth Models and the Evolution of Prolonged Growth Exemplified by Human Brain Evolution.\u201d <em>Journal of Theoretical Biology<\/em> 190 (3): 227\u2013239.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Vrba, Elisabeth S. 2000. \u201cMajor Features of Neogene Mammalian Evolution in Africa.\u201d In <em>Cenozoic <\/em><em>G<\/em><em>eology of <\/em><em>S<\/em><em>outhern Africa<\/em>, edited by T. C. Partridge and R. Maud, 277\u2013304<em>.<\/em> Oxford: Oxford University Press.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Walker, Alan C., and Richard E. Leakey. 1988. \u201cThe Evolution of <em>Australopithecus boisei<\/em>.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by F. E. Grine, 247\u2013258. New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Walker, Alan, Richard E. Leakey, John M. Harris, and Francis H. Brown. 1986. \u201c2.5-my <em>Australopithecus boisei<\/em> from West of Lake Turkana, Kenya.\u201d <em>Nature<\/em> 322 (6079): 517\u2013522.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Ward, Carol, Meave Leakey, and Alan Walker. 1999. \u201cThe New Hominid Species <em>Australopithecus anamensis<\/em>.\u201d <em>Evolutionary Anthropology<\/em> 7 (6): 197\u2013205.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">White, Tim D. 1988. \u201cThe Comparative Biology of \u2018Robust\u2019 Australopithecus: Clues from Content.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by F. E. Grine, 449\u2013483. New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">White, Tim D., Gen Suwa, and Berhane Asfaw. 1994. \u201c<em>Australopithecus ramidus<\/em>, a New Species of Early Hominid from Aramis, Ethiopia.\u201d <em>Nature<\/em> 371 (6495): 306\u2013312.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Wood, Bernard. 2010. \u201cReconstructing Human Evolution: Achievements, Challenges, and Opportunities.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 10 (2): 8902\u20138909.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Wood, Bernard, and Eve K. Boyle. 2016. \u201cHominin Taxic Diversity: Fact or Fantasy?\u201d <em>Yearbook of Physical Anthropology<\/em> 159 (S61): 37\u201378.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Wood, Bernard, and Kes Schroer. 2017. \u201cParanthropus: Where Do Things Stand?\u201d In <em>Human Paleontology and Prehistory<\/em>, edited by A. Marom and E. Hovers, 95\u2013107. New York: Springer, Cham.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Acknowledgements<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">All of the authors in this section are students and early career researchers in paleoanthropology and related fields in South Africa (or at least have worked in South Africa). We wish to thank everyone who supports young and diverse talent in this field and would love to further acknowledge Black, African, and female academics who have helped pave the way for us.<\/span><\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1767\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1767\"><div tabindex=\"-1\"><p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kerryn Warren, Ph.D., Grad Coach International<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lindsay Hunter, M.A., University of Iowa<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Navashni Naidoo, M.Sc., University of Cape Town<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Silindokuhle Mavuso, M.Sc., University of Witwatersrand<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\"><em>This chapter is a revision from <\/em><em>\"<\/em><a class=\"rId7\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\"><em>Chapter 9: Early Hominins<\/em><\/a><em>\" <\/em><em>by Kerryn Warren, K. Lindsay Hunter, Navashni Naidoo, Silindokuhle Mavuso, Kimberleigh Tommy, Rosa Moll, and Nomawethu Hlazo<\/em><em>. In <\/em><a class=\"rId8\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, first edition<\/em><\/a><em>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under <\/em><a class=\"rId9\" style=\"color: #000000\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\"><em>CC BY-NC 4.0<\/em><\/a><em>. <\/em><\/span><\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\"><span style=\"color: #000000\"><span style=\"color: #ffffff\">Learning Objectives<\/span><br \/>\n<\/span><\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li><span style=\"color: #000000\">Understand what is meant by \u201cderived\u201d and \u201cancestral\u201d traits and why this is relevant for understanding early hominin evolution.<\/span><\/li>\n<li><span style=\"color: #000000\">Understand changing paleoclimates and paleoenvironments as potential factors influencing early hominin adaptations.<\/span><\/li>\n<li><span style=\"color: #000000\">Describe the anatomical changes associated with bipedalism and dentition in early hominins, as well as their implications..<\/span><\/li>\n<li><span style=\"color: #000000\">Describe early hominin genera and species, including their currently understood dates and geographic expanses.<\/span><\/li>\n<li><span style=\"color: #000000\">Describe the earliest stone tool techno-complexes and their impact on the transition from early hominins to our genus.<\/span><\/li>\n<\/ul>\n<\/div>\n<\/div>\n<h2 class=\"__UNKNOWN__\"><span style=\"color: #000000\">Defining Hominins<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">It is through our study of our hominin ancestors and relatives that we are exposed to a world of \u201cmight have beens\u201d: of other paths not taken by our species, other ways of being human. But to better understand these different evolutionary trajectories, we must first define the terms we are using. If an imaginary line were drawn between ourselves and our closest relatives, the great apes, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1200\">bipedalism<\/a><\/strong> (or habitually walking upright on two feet) is where that line would be. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1270\">Hominin<\/a><\/strong>, then, means everyone on \u201cour\u201d side of the line: humans and all of our extinct bipedal ancestors and relatives since our divergence from the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1272\">last common ancestor (LCA)<\/a><\/strong> we share with chimpanzees.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Historic interpretations of our evolution, prior to our finding of early hominin <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1274\"><strong>fossils<\/strong><\/a>, varied. Debates in the mid-1800s regarding hominin origins focused on two key issues:<\/span><\/p>\n<ul>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">Where did we evolve?<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">Which traits evolved first?<\/span><\/li>\n<\/ul>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff00ff\">Charles Darwin hypothesized that we evolved in Africa, as he was convinced that we shared greater commonality with chimpanzees and gorillas on the continent (Darwin 1871). Others, such as Ernst Haeckel and Eug\u00e8ne Dubois, insisted that we were closer in affinity to orangutans and that we evolved in Eurasia where, until the discovery of the Taung Child in South Africa in 1924, all humanlike fossils (of Neanderthals and <em>Homo erectus<\/em>) had been found (Shipman 2002).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Within this conversation, naturalists and early <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1276\">paleoanthropologists<\/a><\/strong> (people who study human evolution) speculated about which human traits came first. These included the evolution of a big brain (<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1278\"><strong>encephalization<\/strong><\/a>), the evolution of the way in which we move about on two legs (bipedalism), and the evolution of our flat faces and small teeth (indications of dietary change). Original hypotheses suggested that, in order to be motivated to change diet and move about in a bipedal fashion, the large brain needed to have evolved first, as is seen in the fossil species mentioned above.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">However, we now know that bipedal locomotion is one of the first things that evolved in our lineage, with early relatives having more apelike dentition and small brain sizes. While brain size expansion is seen primarily in our genus, <em>Homo<\/em>, earlier hominin brain sizes were highly variable between and within taxa, from 300 cc (cranial capacity, cm<sup>3<\/sup>), estimated in <em>Ardipithecus<\/em>, to 550 cc, estimated in <em>Paranthropus boisei<\/em>. The lower estimates are well within the range of variation of nonhuman extant great apes. In addition, body size variability also plays a role in the interpretation of whether brain size could be considered large or small for a particular species or specimen. In this chapter, we will tease out the details of early hominin evolution in terms of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1280\"><strong>morphology<\/strong><\/a> (i.e. the study of the form, size, or shape of things; in this case, skeletal parts).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">We also know that early human evolution occurred in a very complicated fashion. There were multiple species (multiple genera) that featured diversity in their diets and locomotion. Specimens have been found all along the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1282\"><strong>East African Rift System <\/strong>(<strong>EARS)<\/strong><\/a>; that is, in Ethiopia, Kenya, Tanzania, and Malawi; see Figure 9.1), in limestone caves in South Africa, and in Chad. Dates of these early relatives range from around 7 million years ago (mya) to around 1 mya, overlapping temporally with members of our genus, <em>Homo<\/em>.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 610px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/06\/image38.png\" alt=\"Patchy green mountain alongside a deep sandy valley in East Africa.\" width=\"610\" height=\"277\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.1: East African Rift System (EARS). Credit: <a href=\"https:\/\/www.flickr.com\/photos\/ninara\/8624605781\/in\/photolist-x2yH7-x2yHe-VfVWuD-e98mPF-SzzjsU-2bsBZhC-2hHec7m-xtJ7Ez-NXnXvh-7Yg3uo-2cS3FgG-2hjo1Dc-2hjGoTS-nnumi8-82U66W-dMNn7B-8jdVbd-NWDg8-NW6fj-ebhx5w-bkFv1G-Ct5ZD-5JQk8A-y6TgAc-x9k6oe-2ebLTDC-WcPMnJ-2ekh6CS-Cu3LH-xNHDFK-9RUsZi-94jVt4-P46uiB-QFyjyE-crU8N7-5JLJKV-2ekSgk8-5JL454-2cPgZrF-2bHfQZu-dMTVPN-6yUbeN-jzMicQ-48XjU9-2etR2Ze-Styrvw-crU7V7-2wakq3-crU6Z1-2etR2XR\/\">IMG_1696 Great Rift Valley<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/ninara\/\">Ninara<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\">CC BY 2.0 License<\/a>.<\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Yet there is still so much to understand. Modern debates now look at the relatedness of these species to us and to one another, and they consider which of these species were able to make and use tools. As a result, every <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1284\">site<\/a><\/strong> discovery in the patchy hominin fossil record tells us more about our evolution. In addition, recent scientific techniques (not available even ten years ago) provide new insights into the diets, environments, and lifestyles of these ancient relatives.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In the past, <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1288\"><strong>taxonom<\/strong><strong>y<\/strong><\/a> was primarily based on morphology. Today it is tied to known relationships based on molecular <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1290\"><strong>phylogeny<\/strong><\/a> (e.g., based on DNA) or a combination of the two. This is complicated when applied to living <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1286\"><strong>taxa<\/strong><\/a>, but becomes much more difficult when we try to categorize ancestor-descendant relationships for long-extinct species whose molecular information is no longer preserved. We therefore find ourselves falling back on morphological comparisons, often of teeth and partially fossilized skeletal material.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">It is here that we turn to the related concepts of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1292\">cladistics<\/a> <\/strong>and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1294\">phylogenetics<\/a><\/strong><strong>. <\/strong><span style=\"background-color: #ff99cc\">Cladistics groups organisms according to their last common ancestors based on shared <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1126\">derived traits<\/a><\/strong>. <\/span>In the case of early hominins, these are often morphological traits that differ from those seen in earlier populations. These new or modified traits provide evidence of evolutionary relationships, and organisms with the same derived traits are grouped in the same <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1116\">clade<\/a> <\/strong>(Figure 9.2). <span style=\"background-color: #ff99cc\">For example, if we use feathers as a trait, we can group pigeons and ostriches into the clade of birds.<\/span> In this chapter, we will examine the grouping of the Robust Australopithecines, whose cranial and dental features differ from those of earlier hominins, and therefore are considered derived.<\/span><\/p>\n<figure style=\"width: 708px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image48.png\" alt=\"Phylogenetic tree shows clades and non clade groupings.\" width=\"708\" height=\"192\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.2: Clades refer to groups of species or taxa that share a common ancestor. In <span class=\"ILfuVd\" lang=\"en\"><span class=\"hgKElc\">a phylogeny, a clade is a complete group of lineages, including their last common ancestor. Groupings that do not include a common ancestor and <em>all<\/em> of its descendants are not clades. <\/span><\/span>Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\">Clades (Figure 9.2)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Katie Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Dig Deeper: Problems Defining Hominin Species<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000\">It is worth noting that species designations for early hominin specimens are often highly contested. This is due to the fragmentary nature of the fossil record, the large timescale (millions of years) with which paleoanthropologists need to work, and the difficulty in evaluating whether morphological differences and similarities are due to meaningful phylogenetic or biological differences or subtle differences\/variation in niche occupation or time. In other words, do morphological differences really indicate different species? How would classifying species in the paleoanthropological record compare with classifying living species today, for whom we can sequence genomes and observe lifestyles?<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">There are also broader philosophical differences among researchers when it comes to paleo-species designations. Some scientists, known as \u201c<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1296\"><strong>lumpers<\/strong>,<\/a>\u201d argue that large variability is expected among multiple populations in a given species over time. These researchers will therefore prefer to \u201clump\u201d specimens of subtle differences into single taxa. Others, known as \u201c<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1298\"><strong>splitters<\/strong><\/a>,\u201d argue that species variability can be measured and that even subtle differences can imply differences in niche occupation that are extreme enough to mirror modern species differences. In general, splitters would consider geographic differences among populations as meaning that a species is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1300\">polytypic<\/a><\/strong> <span style=\"background-color: #ff99cc\">(i.e., capable of interacting and breeding biologically but having morphological population differences)<\/span>. This is worth keeping in mind when learning about why species designations may be contested.<\/span><\/p>\n<figure style=\"width: 520px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image8.jpg\" alt=\"A graph shows a curved line depicting changes in morphology among two species over time.\" width=\"520\" height=\"292\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.3: This graph demonstrates the concept of a chronospecies, where one species (Species A) \u201cevolves\u201d into another (Species B). Credit: Chronospecies original to Explorations: An Open Invitation to Biological Anthropology, 2nd edition by Kerryn Warren is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\"><span style=\"color: #000000\">This further plays a role in evaluating ancestry. Debates over which species \u201cgave rise\u201d to which continue to this day. It is common to try to create \u201clineages\u201d of species to determine when one species evolved into another over time. We refer to these as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1302\"><strong>chronospecies<\/strong><\/a> (Figure 9.3). Constructed hominin phylogenetic trees are routinely variable, changing with new specimen discoveries, new techniques for evaluating and comparing species, and, some have argued, nationalist or biased interpretations of the record. More recently, some researchers have shifted away from \u201ctreelike\u201d models of ancestry toward more nuanced metaphors such as the \u201cbraided stream,\u201d where some levels of interbreeding among species and populations are seen as natural processes of evolution.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Finally, it is worth considering the process of fossil discovery and publication. Some fossils are easily diagnostic to a species level and allow for easy and accurate interpretation. Some, however, are more controversial. This could be because they do not easily preserve or are incomplete, making it difficult to compare and place within a specific species (e.g., a fossil of a patella or knee bone). Researchers often need to make several important claims when announcing or publishing a find: a secure date (if possible), clear association with other finds, and an adequate comparison among multiple species (both extant and fossil). Therefore, it is not uncommon that an important find was made years before it is scientifically published.<\/span><\/p>\n<\/div>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Paleoenvironment and Hominin Evolution<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There is no doubt that one of the major selective pressures in hominin evolution is the environment. <span style=\"background-color: #ff9900\"><span style=\"background-color: #ffff00\">Large-scale changes in global and regional climate, as well as alterations to the environment, are<\/span><span style=\"background-color: #ffff00\"> (<\/span><\/span><em style=\"background-color: transparent\"><span style=\"text-decoration: underline\">thought to be<\/span>)<\/em><span style=\"background-color: #ffff00\"><del> all<\/del> linked to (all) hominin diversification, dispersal, and extinction<\/span> (Maslin et al. 2014). Environmental reconstructions often use modern analogues. Let us take, for instance, the hippopotamus. It is an animal that thrives in environments that have abundant water to keep its skin cool and moist. If the environment for some reason becomes drier, it is expected that hippopotamus populations will reduce. If a drier environment becomes wetter, it is possible that hippopotamus populations may be attracted to the new environment and thrive. Such instances have occurred multiple times in the past, and the bones of some <strong style=\"background-color: transparent\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1308\">fauna<\/a><\/strong> (i.e., animals, like the hippopotamus) that are sensitive to these changes give us insights into these events.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Yet reconstructing a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1304\">paleoenvironment<\/a><\/strong> relies on a range of techniques, which vary depending on whether research interests focus on local changes or more global environmental changes\/reconstructions. For local environments (such as a single site or region), comparing the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1306\">faunal assemblages<\/a> <\/strong>(collections of fossils of animals found at a site) with animals found in certain modern environments allows us to determine if past environments mirror current ones in the region. Changes in the faunal assemblages, as well as when they occur and how they occur, tell us about past environmental changes. Other techniques are also useful in this regard. Chemical analyses, for instance, can reveal the diets of individual fauna, providing clues as to the relative wetness or dryness of their environment (e.g., nitrogen <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1310\"><strong>isotopes<\/strong><\/a>; Kingston and Harrison 2007).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Global climatic changes in the distant past, which fluctuated between being colder and drier and warmer and wetter on average, would have global implications for environmental change (Figure 9.4). These can be studied by comparing marine core and terrestrial soil data across multiple sites. These techniques are based on chemical analysis, such as examination of the nitrogen and oxygen isotopes in shells and sediments. Similarly, analyzing pollen grains shows which kinds of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1312\"><strong>flora<\/strong> <\/a> survived in an environment at a specific time period. There are multiple lines of evidence that allow us to visualize global climate trends over millions of years (although it should be noted that the direction and extent of these changes could differ by geographic region).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 649px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image12-1-1.png\" alt=\"Chart shows cyclical carbon dioxide levels from 800,000 years ago until today.\" width=\"649\" height=\"406\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.4: This graph, based on the comparison of atmospheric samples contained in ice cores and more recent direct measurements, illustrates how atmospheric CO\u2082 has fluctuated over time and increased sharply since the Industrial Revolution. The graph also shows that since 800,000ya (and before) atmospheric CO\u2082 has never exceeded 300 parts per million (ppm). In 1950 it was 310ppm. Today atmospheric CO\u2082 has spiked to over 410 ppm. Credit: <a href=\"https:\/\/climate.nasa.gov\/evidence\/\">CO\u2082 increase since the Industrial Revolution<\/a> by <a href=\"https:\/\/www.nasa.gov\/\">NASA<\/a> is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a> and is used within <a href=\"https:\/\/www.nasa.gov\/multimedia\/guidelines\/index.html\">NASA guidelines on re-use<\/a>. Original from Luthi, D., et al.. 2008; Etheridge, D.M., et al. 2010; Vostok ice core data\/J.R. Petit et al.; NOAA Mauna Loa CO<a href=\"https:\/\/climate.nasa.gov\/evidence\/\">\u2082<\/a> record..<\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Both local and global climatic\/environmental changes have been used to understand factors affecting our evolution (DeHeinzelin et al. 1999; Kingston 2007). Environmental change acts as an important factor regarding the onset of several important hominin traits seen in early hominins and discussed in this chapter. Namely, the environment has been interpreted as the following:<\/span><\/p>\n<ul>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">the driving force behind the evolution of bipedalism,<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">the reason for change and variation in early hominin diets, and<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">the diversification of multiple early hominin species.<\/span><\/li>\n<\/ul>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are numerous hypotheses regarding how climate has driven and continues to drive human evolution. Here, we will focus on just three popular hypotheses.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Savannah Hypothesis (or Aridity Hypothesis)<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The hypothesis:<\/strong> This popular theory suggests that the expansion of the savannah (or less densely forested, drier environments) forced early hominins from an <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1314\"><strong>arboreal<\/strong> <\/a> lifestyle (one living in trees) to a terrestrial one where bipedalism was a more efficient form of locomotion (Figure 9.5). It was first proposed by Darwin (1871) and supported by anthropologists like Raymond Dart (1925). However, this idea was supported by little fossil or paleoenvironmental evidence and was later refined as the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1318\">Aridity Hypothesis<\/a><\/strong>. This hypothesis states that the long-term <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1316\">aridification<\/a><\/strong> and, thereby, expansion of savannah biomes were drivers in diversification in early hominin evolution (deMenocal 2004; deMenocal and Bloemendal 1995). It advocates for periods of accelerated aridification leading to early hominin speciation events.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 647px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image46.png\" alt=\"Photograph showing a dry, open savannah environment.\" width=\"647\" height=\"486\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.5: The African savannah grew during early hominin evolution. This may have forced early hominins from an arboreal lifestyle to a terrestrial one, where bipedalism was a more efficient form of locomotion. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:African_savannah_@_Masai_Mara_(21308330314).jpg\">African savannah @ Masai Mara (21308330314)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/132394214@N04\">Leo Li<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\">CC BY 2.0 License<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The evidence:<\/strong> While early bipedal hominins are often associated with wetter, more closed environments (i.e., not the Savannah Hypothesis), both marine and terrestrial records seem to support general cooling, drying conditions, with isotopic records indicating an increase in grasslands (i.e., colder and wetter climatic conditions) between 8 mya and 6 mya across the African continent (Cerling et al. 2011). This can be contrasted with later climatic changes derived from aeolian dust records (sediments transported to the site of interest by wind), which demonstrate increases in seasonal rainfall between 3 mya and 2.6 mya, 1.8 mya and 1.6 mya, and 1.2 mya and 0.8 mya (deMenocal 2004; deMenocal and Bloemendal 1995).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Interpretation(s):<\/strong> Despite a relatively scarce early hominin record, it is clear that two important factors occur around the time period in which we see increasing aridity. The first factor is the diversification of taxa, where high morphological variation between specimens has led to the naming of multiple hominin genera and species. The second factor is the observation that the earliest hominin fossils appear to have traits associated with bipedalism and are dated to around the drying period (as based on isotopic records). Some have argued that it is more accurately a combination of bipedalism and arboreal locomotion, which will be discussed later. However, the local environments in which these early specimens are found (as based on the faunal assemblages) do not appear to have been dry.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Turnover Pulse Hypothesis<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The hypothesis:<\/strong> In 1985, paleontologist Elisabeth Vbra noticed that in periods of extreme and rapid climate change, <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1326\"><strong>ungulates<\/strong><\/a> (hoofed mammals of various kinds) that had generalized diets fared better than those with specialized diets (Vrba 1988, 1998). <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1322\"><strong>Specialist<\/strong><\/a> eaters <span style=\"background-color: #ff99cc\">(those who rely primarily on specific food types)<\/span> faced extinction at greater rates than their <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1320\">generalist<\/a> <\/strong><span style=\"background-color: #ff99cc\">(those who can eat more varied and variable diets) <\/span>counterparts because they were unable to adapt to new environments (Vrba 2000). Thus, periods with extreme climate change would be associated with high <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1324\"><strong>faunal turnover<\/strong><\/a>: that is, the extinction of many species and the speciation, diversification, and migration of many others to occupy various niches.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The evidence:<\/strong> The onset of the<strong> <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1400\">Quaternary Ice Age<\/a><\/strong>, between 2.5 mya and 3 mya, brought extreme global, cyclical <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1328\"><strong>interglacial<\/strong> <\/a> and <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1330\"><strong>glacial<\/strong><\/a> periods (warmer, wetter periods with less ice at the poles, and colder, drier periods with more ice near the poles). Faunal evidence from the Turkana basin in East Africa indicates multiple instances of faunal turnover and extinction events, in which global climatic change resulted in changes from closed\/forested to open\/grassier habitats at single sites (Behrensmeyer et al. 1997; Bobe and Behrensmeyer 2004). Similarly, work in the Cape Floristic Belt of South Africa shows that extreme changes in climate play a role in extinction and migration in ungulates. While this theory was originally developed for ungulates, its proponents have argued that it can be applied to hominins as well. However, the link between climate and speciation is only vaguely understood (Faith and Behrensmeyer 2013).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Interpretation(s):<\/strong> While the evidence of rapid faunal turnover among ungulates during this time period appears clear, there is still some debate around its usefulness as applied to the paleoanthropological record. Specialist hominin species do appear to exist for long periods of time during this time period, yet it is also true that <em>Homo<\/em>, a generalist genus with a varied and adaptable diet, ultimately survives the majority of these fluctuations, and the specialists appear to go extinct.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Variability Selection Hypothesis<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The hypothesis: <\/strong>This hypothesis was first articulated by paleoanthropologist Richard Potts (1998). It links the high amount of climatic variability over the last 7 million years to both behavioral and morphological changes. Unlike previous notions, this hypothesis states that hominin evolution does not respond to habitat-specific changes or to specific aridity or moisture trends. Instead, long-term environmental unpredictability over time and space influenced morphological and behavioral adaptations that would help hominins survive, regardless of environmental context (Potts 1998, 2013). The Variability Selection Hypothesis states that hominin groups would experience varying degrees of natural selection due to continually changing environments and potential group isolation. This would allow certain groups to develop genetic combinations that would increase their ability to survive in shifting environments. These populations would then have a genetic advantage over others that were forced into habitat-specific adaptations (Potts 2013).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The evidence:<\/strong> The evidence for this theory is similar to that for the Turnover Pulse Hypothesis: large climatic variability and higher survivability of generalists versus specialists. However, this hypothesis accommodates for larger time-scales of extinction and survival events.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Interpretation(s):<\/strong> In this way, the Variability Selection Hypothesis allows for a more flexible interpretation of the evolution of bipedalism in hominins and a more fluid interpretation of the Turnover Pulse Hypothesis, where species turnover is meant to be more rapid. In some ways, this hypothesis accommodates both environmental data and our interpretations of an evolution toward greater variability among species and the survivability of generalists.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000;background-color: #ff99cc\"><strong>Paleoenvironment Summary<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff99cc\">Some hypotheses presented in this section pay specific attention to habitat (Savannah Hypothesis) while others point to large-scale climatic forces (Variability Selection Hypothesis). Some may be interpreted to describe the evolution of traits such as bipedalism (Savannah Hypothesis), and others generally explain the diversification of early hominins (Turnover Pulse and Variability Selection Hypotheses). While there is no consensus as to how the environment drove our evolution, it is clear that the environment shaped both habitat and resource availability in ways that would have influenced our early ancestors physically and behaviorally.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Derived Adaptations: Bipedalism<br \/>\n<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The unique form of locomotion exhibited by modern humans, called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1332\">obligate bipedalism<\/a><\/strong>, is important in distinguishing our species from the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1334\"><strong>extant<\/strong><\/a> (living) great apes. The ability to walk habitually upright is thus considered one of the defining attributes of the hominin lineage. We also differ from other animals that walk bipedally (such as kangaroos) in that we do not have a tail to balance us as we move.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The origin of bipedalism in hominins has been debated in paleoanthropology, but at present there are two main <span style=\"background-color: #ff99cc\">ideas<\/span>: <span style=\"text-decoration: underline\">(<em>theories<\/em>)<\/span><\/span><\/p>\n<ol>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">early hominins initially lived in trees, but increasingly started living on the ground, so we were a product of an arboreal last common ancestor (LCA) or,<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">our LCA was a terrestrial quadrupedal knuckle-walking species, more similar to extant chimpanzees.<\/span><\/li>\n<\/ol>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Most research supports the first theory of an arboreal LCA based on skeletal morphology of early hominin genera that demonstrate adaptations for climbing but not for knuckle-walking. This would mean that both humans and chimpanzees can be considered \u201cderived\u201d in terms of locomotion since chimpanzees would have independently evolved knuckle-walking.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are many current ideas regarding selective pressures that would lead to early hominins adapting upright posture and locomotion. Many of these selective pressures, as we have seen in the previous section, coincide with a shift in environmental conditions, supported by paleoenvironmental data. In general, however, it appears that, like extant great apes, early hominins thrived in forested regions with dense tree coverage, which would indicate an arboreal lifestyle. As the environmental conditions changed and a savannah\/grassland environment became more widespread, the tree cover would become less dense, scattered, and sparse such that bipedalism would become more important.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are several proposed selective pressures for bipedalism:<\/span><\/p>\n<ol>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\"><strong>E<\/strong><strong>nergy conservation:<\/strong> Modern bipedal humans conserve more energy than extant chimpanzees, which are predominantly knuckle-walking quadrupeds when walking over land. While chimpanzees, for instance, are faster than humans terrestrially, they expend large amounts of energy being so. Adaptations to bipedalism include \u201cstacking\u201d the majority of the weight of the body over a small area around the center of gravity (i.e., the head is above the chest, which is above the pelvis, which is over the knees, which are above the feet). This reduces the amount of muscle needed to be engaged during locomotion to \u201cpull us up\u201d and allows us to travel longer distances expending far less energy.<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\"><strong>T<\/strong><strong>hermoregulation:<\/strong> Less surface area (i.e., only the head and shoulders) is exposed to direct sunlight during the hottest parts of the day (i.e., midday). This means that the body has less need to employ additional \u201ccooling\u201d mechanisms such as sweating, which additionally means less water loss.<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Bipedalism <\/strong><span style=\"text-decoration: underline\">(Freeing of Hands)<\/span><strong>: <\/strong>This method of locomotion freed up our ancestors\u2019 hands such that they could more easily gather food and carry tools or infants. This further enabled the use of hands for more specialized adaptations associated with the manufacturing and use of tools.<\/span><\/li>\n<\/ol>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">These selective pressures are not mutually exclusive. Bipedality could have evolved from a combination of these selective pressures, in ways that increased the chances of early hominin survival.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Skeletal Adaptations for Bipedalism<\/strong><\/span><\/h3>\n<figure style=\"width: 405px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image10-1.png\" alt=\"A full human skeleton and gorilla skeleton standing in upright positions next to each other.\" width=\"405\" height=\"452\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.6: Compared to gorillas (right) and other apes, humans (left) have highly specialized adaptations to facilitate bipedal locomotion. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Primatenskelett-drawing.jpg\">Skeleton of human (1) and gorilla (2), unnaturally sketched<\/a> by unknown from Brehms Tierleben, Small Edition 1927 is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Humans have highly specialized adaptations to facilitate obligate bipedalism (Figure 9.6). Many of these adaptations occur within the soft tissue of the body (e.g., muscles and tendons). However, when analyzing the paleoanthropological record for evidence of the emergence of bipedalism, all that remains is the fossilized bone. Interpretations of locomotion are therefore often based on comparative analyses between fossil remains and the skeletons of extant primates with known locomotor behaviors. These adaptations occur throughout the skeleton and are summarized in Figure 9.7.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The majority of these adaptations occur in the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1336\"><strong>postcranium<\/strong><\/a> <span style=\"background-color: #ff99cc\">(the skeleton from below the head)<\/span> and are outlined in Figure 9.7. In general, these adaptations allow for greater stability and strength in the lower limb, by allowing for more shock absorption, for a larger surface area for muscle attachment, and for the \u201cstacking\u201d of the skeleton directly over the center of gravity to reduce energy needed to be kept upright. These adaptations often mean less flexibility in areas such as the knee and foot.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">However, these adaptations come at a cost. Evolving from a nonobligate bipedal ancestor means that the adaptations we have are evolutionary compromises. For instance, the valgus knee (angle at the knee) is an essential adaptation to balance the body weight above the ankle during bipedal locomotion. However, the strain and shock absorption at an angled knee eventually takes its toll. For example, runners often experience joint pain. Similarly, the long neck of the femur absorbs stress and accommodates for a larger pelvis, but it is a weak point, resulting in hip replacements being commonplace among the elderly, especially in cases where the bone additionally weakens through osteoporosis. Finally, the S-shaped curve in our spine allows us to stand upright, relative to the more curved C-shaped spine of an LCA. Yet the weaknesses in the curves can lead to pinching of nerves and back pain. Since many of these problems primarily are only seen in old age, they can potentially be seen as an evolutionary compromise.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Despite relatively few postcranial fragments, the fossil record in early hominins indicates a complex pattern of emergence of bipedalism. Key features, such as a more anteriorly placed foramen magnum, are argued to be seen even in the earliest discovered hominins, indicating an upright posture (Dart 1925). Some early species appear to have a mix of ancestral (arboreal) and derived (bipedal) traits, which indicates a mixed locomotion and a more <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1402\">mosaic evolution<\/a><\/strong> of the trait. Some early hominins appear to, for instance, have bowl-shaped pelvises (hip bones) and angled femurs suitable for bipedalism but also have retained an opposable <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1338\">hallux<\/a><\/strong> (big toe) or curved fingers and longer arms (for arboreal locomotion). These mixed morphologies may indicate that earlier hominins were not fully obligate bipeds and thus thrived in mosaic environments.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Yet the associations between postcranial and the more diagnostic cranial fossils and bones are not always clear, muddying our understanding of the specific species to which fossils belong (Grine et al. 2022).<\/span><\/p>\n<div align=\"left\">\n<table class=\"grid\">\n<caption>\n<p class=\"import-Normal\" style=\"text-align: left\"><span style=\"color: #000000\">Figure 9.7: Skeletal comparisons between modern humans (obligate bipeds) and nonobligate bipeds (e.g., chimpanzees). Credit: <a class=\"rId34\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\">Skeletal comparisons between modern humans and <\/a><a class=\"rId35\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\">nonobligate<\/a><a class=\"rId36\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\"> bipeds (Figure 9.6)<\/a> original to <a class=\"rId37\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a class=\"rId38\" style=\"color: #000000\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/span><\/p>\n<\/caption>\n<thead>\n<tr>\n<td style=\"width: 97.998px\"><strong>Region<\/strong><\/td>\n<td style=\"width: 106.992px\"><strong>Feature<\/strong><\/td>\n<td style=\"width: 366.992px\"><strong>Obligate Biped (H. sapiens)<\/strong><\/td>\n<td style=\"width: 310px\"><strong>Nonobligate Biped<\/strong><\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr>\n<td style=\"width: 97.998px\">Cranium<\/td>\n<td style=\"width: 106.992px\">Position of the foramen magnum<\/td>\n<td style=\"width: 366.992px\">Positioned inferiorly (immediately under the cranium) so that the head rests on top of the vertebral column for balance and support (head is perpendicular to the ground).<\/td>\n<td style=\"width: 310px\">Posteriorly positioned (to the back of the cranium). Head is positioned parallel to the ground.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Body proportions<\/td>\n<td style=\"width: 366.992px\">Shorter upper limb (not used for locomotion).<\/td>\n<td style=\"width: 310px\">Longer upper limbs (used for locomotion).<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Spinal curvature<\/td>\n<td style=\"width: 366.992px\">S-curve due to pressure exerted on the spine from bipedalism (lumbar lordosis).<\/td>\n<td style=\"width: 310px\">C-curve.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Vertebrae<\/td>\n<td style=\"width: 366.992px\">Robust lumbar (lower-back) vertebrae (for shock absorbance and weight bearing). Lower back is more flexible than that of apes as the hips and trunk swivel when walking (weight transmission).<\/td>\n<td style=\"width: 310px\">Gracile lumbar vertebrae compared to those of modern humans.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Pelvis<\/td>\n<td style=\"width: 366.992px\">Shorter, broader, bowl-shaped pelvis (for support); very robust. Broad sacrum with large sacroiliac joint surfaces.<\/td>\n<td style=\"width: 310px\">Longer, flatter, elongated ilia; more narrow and gracile; narrower sacrum; relatively smaller sacroiliac joint surface.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Lower limb<\/td>\n<td style=\"width: 366.992px\">In general, longer, more robust lower limbs and more stable, larger joints.<\/p>\n<ul>\n<li style=\"font-weight: 400\">Large femoral head and longer neck (absorbs more stress and increases the mechanical advantage).<\/li>\n<li style=\"font-weight: 400\">Valgus knee, in which the angle of the knee positions it over the ankle and keeps the center of gravity balanced over the stance leg during stride cycle (shock absorbance).<\/li>\n<li style=\"font-weight: 400\">Distal tibia (lower leg) of humans has a large medial malleolus for stability.<\/li>\n<\/ul>\n<\/td>\n<td style=\"width: 310px\">In general, smaller, more gracile limbs with more flexible joints.<\/p>\n<ul>\n<li style=\"font-weight: 400\">Femoral neck is smaller in comparison to modern humans and shorter.<\/li>\n<li style=\"font-weight: 400\">The legs bow outward, and there is no valgus angle of the knee (no \u201cknock knees\u201d).<\/li>\n<li style=\"font-weight: 400\">The distal tibia in chimpanzees is trapezoid (wider anteriorly) for climbing and allows more flexibility.<\/li>\n<\/ul>\n<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Foot<\/td>\n<td style=\"width: 366.992px\">Rigid, robust foot, without a midtarsal break.<\/p>\n<p>Nonopposable and large, robust big toe (for push off while walking) and large heel for shock absorbance.<\/td>\n<td style=\"width: 310px\">Flexible foot, midtarsal break present (which allows primates to lift their heels independently from their feet), opposable big toe for grasping.<\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<p class=\"import-Normal\"><span style=\"color: #000000;background-color: #ccffcc\">It is also worth noting that, while not directly related to bipedalism per se, other postcranial adaptations are evident in the hominin fossil record from some of the earlier hominins. For instance, the hand and finger morphologies of many of the earliest hominins indicate adaptations consistent with arboreality. These include longer hands, more curved metacarpals and phalanges (long bones in the hand and fingers, respectively), and a shorter, relatively weaker thumb. This allows for gripping onto curved surfaces during locomotion. The earliest hominins appear to have mixed morphologies for both bipedalism and arborealism. However, among Australopiths (members of the genus, Australopithecus), there are indications for greater reliance on bipedalism as the primary form of locomotion. Similarly, adaptations consistent with tool manufacture (shorter fingers and a longer, more robust thumb, in contrast to the features associated with arborealism) have been argued to appear before the genus <em>Homo<\/em>.<\/span><\/p>\n<div class=\"textbox shaded\">\n<h2>Special Topic: Fear of Snakes \u2014 A Cultural or Biological Response?<\/h2>\n<figure style=\"width: 245px\" class=\"wp-caption alignright\"><img src=\"https:\/\/www.researchgate.net\/profile\/David-Penning\/publication\/315305069\/figure\/fig2\/AS:613947958902785@1523388008174\/Constriction-coil-postures-of-kingsnakes-and-ratsnakes-Typical-constriction-coil.png\" alt=\"Constriction coil postures of kingsnakes and ratsnakes. Typical constriction coil postures in a kingsnake, Lampropeltis getula (92 g; A), and a ratsnake, Pantherophis guttatus (86 g; B). Both snakes were constricting similarly sized mice, Mus musculus (12 g). The relative prey mass was 13% for the kingsnake and 13.9% for the ratsnake.\u00a0\" width=\"245\" height=\"245\" \/><figcaption class=\"wp-caption-text\">Constriction coil postures of kingsnakes and ratsnakes. source: https:\/\/www.researchgate.net\/figure\/Constriction-coil-postures-of-kingsnakes-and-ratsnakes-Typical-constriction-coil_fig2_315305069<\/figcaption><\/figure>\n<p>It is suggested that primates have three major predators: raptors, felines, and snakes; however, many studies show that of these carnivores, snakes were one of the first that mammals had to contend with alongside dinosaurs, as felines and raptors evolved at a much slower pace than their reptilian competition. Herpetologists trace the evolution of constricting snakes to about 100 million years ago, and by the time mammals arrived around 75 million years ago, constrictors were\u00a0 already well established as a formidable threat (Greene, 2017). \u00a0Both co-existed for millennia and each sustained selective pressures requiring them to evolve specific traits to survive. When venomous snakes eventually emerged 55 to 65 million years ago, they posed yet an additional threat to proto-primates as they required less distance for the predator to kill (2017). Alongside camouflage and silent movement techniques, it was the development of the snake\u2019s hollow fangs through which to deliver venom that was most transformative to primate evolution. As such, primates evolved their pre-conscious attention, and visual acuity to cope with this new threat; therefore, while snakes were adapting morphologically to feed themselves, they were unwittingly teaching proto-primates valuable lessons in predator detection and reacting appropriately in order to survive.<\/p>\n<p>In a 2009 Harvard University study, Lynne A. Isbell hypothesizes that envenoming snakes are linked to being directly responsible for the origins of the evolving complex brains and superior visual capacity in the lineage of anthropoids leading to humans (Isbell, 2009). Forward-facing eyes for binocular vision, depth perception, enhanced visual acuity, stereoscopic and trichromatic colour vision, all traits necessary for snake detection; and the quick motor responses from the primate\u2019s fight, flight, or freeze defence mechanism to circumvent a snake\u2019s squeeze or bite. Numerous laboratory studies show that humans and primates both sense and visually detect snakes more rapidly than other threatening stimuli (Van Le Et al., 2013). These experiments show that snakes elicited the strongest, fastest responses (Van Le Et al., 2013). This is known as \u2018Snake Detection Theory\u2019 and is the evolution of the primate\u2019s complex brain, visual acuity, and rapid motor responses towards snakes in its environment that are the adaptations needed to live successfully as arboreal beings. It is not fortuitous then, that primates that never coexisted with venomous snakes, such as lemurs in Madagascar, have less visual acuity, better olfaction and smaller brains. Within Isbell\u2019s work, a collaborative study by a group of neuroscientists tested this hypothesis and found that, indeed, there is higher neural firing and activity in multiple areas of the primate brain, notably in the pulvinar, a region\u00a0 responsible for visual attention and oculomotor behaviour (Isbell, L., 2009).<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 509px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/www.worldhistory.org\/img\/r\/p\/750x750\/4941.jpg?v=1747098791\" alt=\"Apophis Defeated\" width=\"509\" height=\"309\" \/><figcaption class=\"wp-caption-text\">Apophis being killed by the knife-wielding cat of Heliopolis, an agent of Re;245 Tomb of Inherkhau, Deir el-Medina (TT 359). Note the very large eye of the snake, the source of its malevolent stare or \u201cevil eye.\u201d source: Weeks, Kent R. (2005) The Illustrated Guide to Luxor: Tombs, Temples and Museums, The American University in Cairo Press, Cairo.<\/figcaption><\/figure>\n<p>Today, the fear of snakes is widespread in humans, often shown through avoidance and disgust. A study in <em data-start=\"197\" data-end=\"244\">The Journal of Ethnobiology and Ethnomedicine<\/em> notes that snakes are over-hunted and excluded from conservation efforts worldwide (Cer\u00edaco, 2012). While cultural factors shape our sentiments, instinct also plays a role\u2014such as the developed avoidance behaviors toward threats like snakes. This blend of instinct and cultural influence is not only seen in behavior but also deeply embedded in the stories we tell. Many cultures depict mythological snakes as harbingers of death or chaos. In the Bible, Satan becomes a snake to tempt Eve. Norse mythology features J\u00f6rmungandr, the world serpent who signals the apocalypse. Egyptian myth tells of Apophis, who battles the sun god Ra nightly. Though sources vary, these myths consistently portray snakes as threats. As such, the widespread fear of snakes may reflect both evolutionary and cultural influences. Understood as an adaptive response inherited from primate ancestors\u2014who developed avoidance behaviors toward potentially dangerous stimuli\u2014and reinforced through myths and religious narratives, the enduring presence of snakes as potent figures of fear across human societies and primate groups highlights the complex intertwining of instinct and cultural meaning in shaping human behavior.<\/p>\n<\/div>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Early Hominins: Sahelanthropus and Orrorin<\/strong><\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">We see evidence for bipedalism in some of the earliest fossil hominins, dated from within our estimates of our divergence from chimpanzees. These hominins, however, also indicate evidence for arboreal locomotion.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The earliest dated hominin find (between 6 mya and 7 mya, based on radiometric dating of volcanic tufts) has been argued to come from Chad and is named <strong><em>Sahelanthropus tchadensis<\/em> <\/strong>(Figure 9.8; Brunet et al. 1995). The initial discovery was made in 2001 by Ahounta Djimdoumalbaye and announced in <em>Nature<\/em> in 2002 by a team led by French paleontologist Michel Brunet. The find has a small cranial capacity (360 cc) and smaller canines than those in extant great apes, though they are larger and pointier than those in humans. <span style=\"background-color: #ffff00\">This implies strongly that, over evolutionary time, the need for display and dominance among males has reduced, as has our sexual dimorphism<\/span>. A short cranial base and a foramen magnum that is more humanlike in positioning have been argued to indicate upright walking.<\/span><\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 640px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-289\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.82.jpg\" alt=\"Four views of a beige-colored skull are seen against a black background.\" width=\"640\" height=\"640\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.8: Sahelanthropus tchadensis exhibits a set of derived features, including a long, low cranium; a small, ape-sized braincase; and relatively reduced prognathism. Credit: aa <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 anterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 posterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 inferior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; and d <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 lateral left view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Initially, the inclusion of <em>Sahelanthropus<\/em> in the hominin family was debated by researchers, since the evidence for bipedalism is based on cranial evidence alone, which is not as convincing as postcranial evidence. Yet, a femur (thigh bone) and ulnae (upper arm bones) thought to belong to <em>Sahelanthropus<\/em> was discovered in 2001 (although not published until 2022). These bones may support the idea that the hominin was in fact a terrestrial biped with arboreal capabilities and behaviors (Daver et al. 2022).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Orrorin tugenensis<\/em><\/strong> (Orrorin meaning \u201coriginal man\u201d), dated to between 6 mya and 5.7 mya, was discovered near Tugen Hills in Kenya in 2000. Smaller <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1340\">cheek teeth<\/a><\/strong> (molars and premolars) than those in even more recent hominins, thick enamel, and reduced, but apelike, canines characterize this species. This is the first species that clearly indicates adaptations for bipedal locomotion, with fragmentary leg, arm, and finger bones having been found but few cranial remains. One of the most important elements discovered was a proximal femur, BAR 1002'00. The femur is the thigh bone, and the proximal part is that which articulates with the pelvis; this is very important for studying posture and locomotion. This femur indicates that <em>Ororrin<\/em> was bipedal, and recent studies suggest that it walked in a similar way to later <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1404\">Pliocene<\/a><\/strong> hominins. Some have argued that features of the finger bones suggest potential tool-making capabilities, although many researchers argue that these features are also consistent with climbing.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Early Hominins: The Genus <em>Ardipithecus<\/em><\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Another genus, <em>Ardipithecus<\/em>, is argued to be represented by at least two species: <em>Ardipithecus (Ar.) ramidus <\/em>and <em>Ar. kadabba<\/em>.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Ardipithecus ramidus<\/em><\/strong> (\u201cramid\u201d means root in the Afar language) is currently the best-known of the earliest hominins (Figure 9.9). Unlike <em>Sahelanthropus<\/em> and<em> Orrorin<\/em>, this species has a large sample size of over 110 specimens from Aramis alone. Dated to 4.4 mya, <em>Ar. ramidus<\/em> was found in Ethiopia (in the Middle Awash region and in Gona). This species was announced in 1994 by American palaeoanthropologist Tim White, based on a partial female skeleton nicknamed \u201cArdi\u201d (ARA-VP-6\/500; White et al. 1994). Ardi demonstrates a mosaic of ancestral and derived characteristics in the postcrania. For instance, she had an opposable big toe (hallux), similar to chimpanzees (i.e., more ancestral), which could have aided in climbing trees effectively. However, the pelvis and hip show that she could walk upright (i.e., it is derived), supporting her hominin status. A small brain (300 cc to 350 cc), midfacial projection, and slight prognathism show retained ancestral cranial features, but the cheek bones are less flared and robust than in later hominins.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 706px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-1310\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.9-scaled-1.jpg\" alt=\"Skull cast and partial skeleton with photographs of some bones and line drawings of others.\" width=\"706\" height=\"453\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.9a and b: Researchers believe that Ardipithecus ramidus was able to walk upright, although not as efficiently as later humans. It possessed the musculature required for tree climbing, and while moving quadrupedally, it likely placed weight on the palms of the hands rather than on the knuckles. Credit: a. <a class=\"rId61\" href=\"https:\/\/boneclones.com\/product\/ardipithecus-ramidus-skull-BH-039\">Ardipithecus ramidus Skull<\/a> by <a class=\"rId62\" href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a> is used by permission and available here under a <a class=\"rId63\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>; b. <a class=\"rId64\" href=\"https:\/\/boneclones.com\/product\/ardipithecus-ramidus-skull-BH-039\">Artist\u2019s rendition of \u201cArdi\u201d skeleton<\/a> by <a class=\"rId65\" href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a> is used by permission and available here under a <a class=\"rId66\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong><em>Ardipithecus kadabba<\/em><\/strong> (the species name means \u201coldest ancestor\u201d in the Afar language) is known from localities on the western margin of the Middle Awash region, the same locality where <em>Ar. ramidus<\/em> has been found. Specimens include mandibular fragments and isolated teeth as well as a few postcranial elements from the Asa Koma (5.5 mya to 5.77 mya) and Kuseralee Members (5.2 mya), well-dated and understood (but temporally separate) volcanic layers in East Africa. This species was discovered in 1997 by paleoanthropologist Dr. Yohannes Haile-Selassie. Originally these specimens were referred to as a subspecies of <em>Ar. ramidus<\/em>. In 2002, six teeth were discovered at Asa Koma and the dental-wear patterns confirmed that this was a distinct species, named <em>Ar. kadabba,<\/em> in 2004. One of the postcranial remains recovered included a 5.2 million-year-old toe bone that demonstrated features that are associated with toeing off (pushing off the ground with the big toe leaving last) during walking, a characteristic unique to bipedal walkers. However, the toe bone was found in the Kuseralee Member, and therefore some doubt has been cast by researchers about its association with the teeth from the Asa Koma Member.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"background-color: #ff99cc\"><strong><span style=\"color: #000000\">Bipedal Trends in Early Hominins: Summary<\/span><\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff99cc\">Trends toward bipedalism are seen in our earliest hominin finds. However, many specimens also indicate retained capabilities for climbing. Trends include a larger, more robust hallux; a more compact foot, with an arch; a robust, long femur, angled at the knee; a robust tibia; a bowl-shaped pelvis; and a more anterior foramen magnum. While the level of bipedality in <em>Salehanthropus<\/em> <em>tchadenisis<\/em> is debated since there are few fossils and no postcranial evidence, <em>Orrorin tugenensis<\/em> and <em>Ardipithecus<\/em> <em>k<\/em><em>adabba <\/em>show clear indications of some of these bipedal trends. However, some retained ancestral traits, such as an opposable hallux in <em>Ardipithecus<\/em>, indicate some retention in climbing ability.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Derived Adaptations: Early Hominin Dention<\/span><\/h2>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">The Importance of Teeth<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Teeth are abundant in the fossil record, primarily because they are already highly mineralized as they are forming, far more so than even bone. Because of this, teeth preserve readily. And, because they preserve readily, they are well-studied and better understood than many skeletal elements. In the sparse hominin (and primate) fossil record, teeth are, in some cases, all we have.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Teeth also reveal a lot about the individual from whom they came. We can tell what they evolved to eat, to which other species they may be closely related, and even, to some extent, the level of sexual dimorphism, or general variability, within a given species. This is powerful information that can be contained in a single tooth. With a little more observation, the wearing patterns on a tooth can tell us about the diet of the individual in the weeks leading up to its death. Furthermore, the way in which a tooth is formed, and the timing of formation, can reveal information about changes in diet (or even mobility) over infancy and childhood, using isotopic analyses. When it comes to our earliest hominin relatives, this information is vital for understanding how they lived.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The purpose of comparing different hominin species is to better understand the functional morphology as it applies to dentition. In this, we mean that the morphology of the teeth or masticatory system (which includes jaws) can reveal something about the way in which they were used and, therefore, the kinds of foods these hominins ate. When comparing the features of hominin groups, it is worth considering modern analogues (i.e., animals with which to compare) to make more appropriate assumptions about diet. In this way, hominin dentition is often compared with that of chimpanzees and gorillas (our close ape relatives), as well as with that of modern humans.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The most divergent group, however, is humans. Humans around the world have incredibly varied diets. Among hunter-gatherers, it can vary from a honey- and plant-rich diet, as seen in the Hadza in Tanzania, to a diet almost entirely reliant on animal fat and protein, as seen in Inuits in polar regions of the world. We are therefore considered generalists, more general than the largely <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1342\"><strong>frugivorous<\/strong><\/a> (fruit-eating) chimpanzee or the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1344\"><strong>folivorous<\/strong><\/a> (foliage-eating) gorilla, as discussed in Chapter 5.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">One way in which all humans are similar is our reliance on the processing of our food. We cut up and tear meat with tools using our hands, instead of using our front teeth (incisors and canines). We smash and grind up hard seeds, instead of crushing them with our hind teeth (molars). This means that, unlike our ape relatives, we can rely more on developing tools to navigate our complex and varied diets. <span style=\"text-decoration: underline\">(We could say)<\/span> Our brain, therefore, is our primary masticatory organ. Evolutionarily, our teeth have reduced in size and our faces are flatter, or more <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1346\">orthognathic<\/a>, <\/strong>partially in response to our increased reliance on our hands and brain to process food. Similarly, a reduction in teeth and a more generalist dental morphology could also indicate an increase in softer and more variable foods, such as the inclusion of more meat. <span style=\"background-color: #ffff00\">These trends begin early on in our evolution.<\/span> The link has been made between some of the earliest evidence for stone tool manufacture, the earliest members of our genus, and the features that we associate with these specimens.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">General Dental Trends in Early Hominins<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Several trends are visible in the dentition of early hominins. However, all tend to have the same <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1348\"><strong>dental formula<\/strong><\/a>. The dental formula tells us how many of each tooth type are present in each quadrant of the mouth. Going from the front of the mouth, this includes the square, flat <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1160\">incisors<\/a><\/strong>; the pointy <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1166\">canines<\/a><\/strong>; the small, flatter <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1350\"><strong>premolars<\/strong><\/a>; and the larger hind <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1164\">molars<\/a><\/strong>. In many primates, from Old World monkeys to great apes, the typical dental formula is 2:1:2:3. This means that if we divide the mouth into quadrants, each has two incisors, one canine, two premolars, and three molars. The eight teeth per quadrant total 32 teeth in all (although some humans have fewer teeth due to the absence of their wisdom teeth, or third molars).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 380px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image24.png\" alt=\"Anterior view of the lower face of a person showing their teeth.\" width=\"380\" height=\"253\" \/><figcaption class=\"wp-caption-text\">Figure 9.10: In humans, our canines are often a similar size to our incisors. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Adult_human_teeth.jpg\">Adult human teeth<\/a> by <a href=\"https:\/\/www.genusfotografen.se\/\">Genusfotografen<\/a> (Tomas Gunnarsson) through <a href=\"https:\/\/wikimedia.se\/\">Wikimedia Sverige<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p><span style=\"color: #000000\">The morphology of the individual teeth is where we see the most change. Among primates, large incisors are associated with food procurement or preparation (such as biting small fruits), while small incisors indicate a diet that may contain small seeds or leaves (where the preparation is primarily in the back of the mouth). Most hominins have relatively large, flat, vertically aligned incisors that <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1396\">occlude<\/a> <\/strong>(touch) relatively well, forming a \u201cbite.\u201d This differs from, for instance, the orangutan, whose teeth stick out (i.e.<em>,<\/em> are <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1352\"><strong>procumbent<\/strong><\/a>).<\/span><\/p>\n<p>&nbsp;<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">While the teeth are often aligned with diet, the canines may be misleading in that regard. We tend to associate pointy, large canines with the ripping required for meat, and the reduction (or, in some animals, the absence) of canines as indicative of herbivorous diets. In humans, our canines are often a similar size to our incisors and therefore considered <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1398\">incisiform<\/a><\/strong> (Figure 9.10). However, our closest relatives all have very long, pointy canines, particularly on their upper dentition. This is true even for the gorilla, which lives almost exclusively on plants. The canines in these instances reveal more about social structure and sexual dimorphism than diet, as large canines often signal <span style=\"background-color: #ffff00\">dominance<\/span>.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Early on in human evolution, we see a reduction in canine size. <em>Sahelanthropus tchadensis<\/em> and <em>Orrorin tugenensis<\/em> both have smaller canines than those in extant great apes, yet the canines are still larger and pointier than those in humans or more recent hominins. <span style=\"background-color: #ffff00\">This implies strongly that, over evolutionary time, the need for display and dominance among males has reduced, as has our sexual dimorphism.<\/span> In <em>Ardipithecus ramidus<\/em>, there is no obvious difference between male and female canine size, yet they are still slightly larger and pointier than in modern humans. This implies a less sexually dimorphic social structure in the earlier hominins relative to modern-day chimpanzees and gorillas.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Along with a reduction in canine size is the reduction or elimination of a canine <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1172\">diastema<\/a>:<\/strong> a gap between the teeth on the mandible that allows room for elongated teeth on the maxilla to \u201cfit\u201d in the mouth. Absence of a diastema is an excellent indication of a reduction in canine size. In animals with large canines (such as baboons), there is also often a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1354\"><strong>honing P3<\/strong><\/a>, where the first premolar (also known as P3 for evolutionary reasons) is triangular in shape, \u201csharpened\u201d by the extended canine from the upper dentition. This is also seen in some early hominins: <em>Ardipithecus<\/em>, for example, has small canines that are almost the same height as its incisors, although still larger than those in recent hominins.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The hind dentition, such as the bicuspid (two cusped) premolars or the much larger molars, are also highly indicative of a generalist diet in hominins. Among the earliest hominins, the molars are larger than we see in our genus, increasing in size to the back of the mouth and angled in such a way from the much smaller anterior dentition as to give these hominins a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1356\"><strong>parabolic<\/strong><\/a> (V-shaped) dental arch. This differs from our living relatives and some early hominins, such as <em>Sahelanthropus<\/em>, whose molars and premolars are relatively parallel between the left and right sides of the mouth, creating a U-shape.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Among more recent early hominins, the molars are larger than those in the earliest hominins and far larger than those in our own genus, <em>Homo.<\/em> Large, short molars with thick <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1358\"><strong>enamel<\/strong><\/a> allowed our early cousins to grind fibrous, coarse foods, such as sedges, which require plenty of chewing. This is further evidenced in the low <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1176\">cusps<\/a>,<\/strong> or ridges, on the teeth, which are ideal for chewing. In our genus, the hind dentition is far smaller than in these early hominins. Our teeth also have medium-size cusps, which allow for both efficient grinding and tearing\/shearing meats.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Understanding the dental morphology has allowed researchers to extrapolate very specific behaviors of early hominins. It is worth noting that while teeth preserve well and are abundant, a slew of other morphological traits additionally provide evidence for many of these hypotheses. Yet there are some traits that are ambiguous. For instance, while there are definitely high levels of sexual dimorphism in <em>Au. afarensis<\/em>, discussed in the next section, the canine teeth are reduced in size, implying that while canines may be useful indicators for sexual dimorphism, it is also worth considering other evidence.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff99cc\">In summary, trends among early hominins include a reduction in procumbency, reduced hind dentition (molars and premolars), a reduction in canine size (more incisiform with a lack of canine diastema and honing P3), flatter molar cusps, and thicker dental enamel. All early hominins have the ancestral dental formula of 2:1:2:3. These trends are all consistent with a generalist diet, incorporating more fibrous foods.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Special Topic: Contested Species<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Many named species are highly debated and argued to have specimens associated with a more variable <em>Au. afarensis <\/em>or <em>Au. anamensis<\/em> species. Sometimes these specimens are dated to times when, or found in places in which, there are \u201cgaps\u201d in the palaeoanthropological record. These are argued to represent chronospecies or variants of <em>Au. afarensis<\/em>. However, it is possible that, with more discoveries, the distinct species types will hold.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><strong><em>Australopithecus bahrelghazali<\/em><\/strong> is dated to within the time period of <em>Au. afarensi<\/em>s (3.6 mya; Brunet et al. 1995) and was the first Australopithecine to be discovered in Chad in central Africa. Researchers argue that the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1360\">holotype<\/a><\/strong>, whom discoverers have named \u201cAbel,\u201d falls under the range of variation of <em>Au. afarensis<\/em> and therefore that <em>A. bahrelghazali<\/em> does not fall into a new species (Lebatard et al. 2008). If \u201cAbel\u201d is a member of <em>Au. afarensis<\/em>, the geographic range of the species would be greatly extended.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">On a different note, <strong><em>Australopithecus <\/em><\/strong><strong><em>deyiremada<\/em><\/strong> (meaning \u201cclose relative\u201d in the Ethiopian language of Afar) is dated to 3.5 mya to 3.3 mya and is based on fossil mandible bones discovered in 2011 in Woranso-Mille (in the Afar region of Ethiopia) by Yohannes Haile-Selassie, an Ethiopian paleoanthropologist (Haile-Selassie et al. 2019). The discovery indicated, in contrast to <em>Au. afarensis<\/em>, smaller teeth with thicker enamel (potentially suggesting a harder diet) as well as a larger mandible and more projecting cheekbones. This find may be evidence that more than one closely related hominin species occupied the same region at the same temporal period (Haile-Selassie et al. 2015; Spoor 2015) or that other <em>Au. afarensis<\/em> specimens have been incorrectly designated. However, others have argued that this species has been prematurely identified and that more evidence is needed before splitting the taxa, since the variation appears subtle and may be due to slightly different niche occupations between populations over time.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><strong><em>Australopithecus garhi<\/em><\/strong> is another species found in the Middle Awash region of Ethiopia. It is currently dated to 2.5 mya (younger than <em>Au. afarensis<\/em>). Researchers have suggested it fills in a much-needed temporal \u201cgap\u201d between hominin finds in the region, with some anatomical differences, such as a relatively large cranial capacity (450 cc) and larger hind dentition than seen in other gracile Australopithecines. Similarly, the species has been argued to have longer hind limbs than <em>Au. afarensis<\/em>, although it was still able to move arboreally (Asfaw et al. 1999). However, this species is not well documented or understood and is based on only several fossil specimens. More astonishingly, crude stone tools resembling Oldowan (which will be described later) have been found in association with <em>Au. garhi<\/em>. While lacking some of the features of the Oldowan, this is one of the earliest technologies found in direct association with a hominin.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><strong><em>Kenyanthopus<\/em><\/strong><strong><em> platyops<\/em><\/strong> (the name \u201cplatyops\u201d refers to its flatter-faced appearance) is a highly contested genus\/species designation of a specimen (KNM-WT 40000) from Lake Turkana in Kenya, discovered by Maeve Leakey in 1999 (Figure 9.11). Dated to between 3.5 mya and 3.2 mya, some have suggested this specimen is an <em>Australopithecus<\/em>, perhaps even <em>Au.<\/em> <em>afarensis<\/em> (with a brain size which is difficult to determine, yet appears small), while still others have placed this specimen in <em>Homo <\/em>(small dentition and flat-orthognathic face). While taxonomic placing of this species is quite divided, the discoverers have argued that this species is ancestral to <em>Homo<\/em>, in particular to <em>Homo <\/em><em>ruldolfensis<\/em> (Leakey et al. 2001). Some researchers have additionally associated the earliest tool finds from Lomekwi, Kenya, temporally (3.3 mya) and in close geographic proximity to this specimen.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 579px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-292 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.11.jpg\" alt=\"Four views of an ancient skull are shown on a black background.\" width=\"579\" height=\"579\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.11: This specimen, KNM WT 40000 (Kenyanthopus platyops), has small detention, a small brain case, and a relatively flat face. Its genus\/species designation remains contested. Credit: a. <a class=\"rId76\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId77\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 anterior view<\/a> by \u00a9<a class=\"rId78\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId79\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId80\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId81\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId82\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 superior view<\/a> by \u00a9<a class=\"rId83\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId84\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId85\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a class=\"rId86\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId87\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 lateral left view<\/a> by \u00a9<a class=\"rId88\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId89\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId90\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; d. <a class=\"rId91\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId92\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 inferior view<\/a> by \u00a9<a class=\"rId93\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId94\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId95\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">The Genus <em>Australopithecus<\/em><br \/>\n<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The Australopithecines are a diverse group of hominins, comprising various species. <em>Australopithecus<\/em> is the given group or genus name. It stems from the Latin word <em>Australo<\/em>, meaning \u201csouthern,\u201d and the Greek word <em>pithecus,<\/em> meaning \u201cape.\u201d Within this section, we will outline these differing species\u2019 geological and temporal distributions across Africa, unique derived and\/or shared traits, and importance in the fossil record.<\/span><\/p>\n<figure style=\"width: 381px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image37-2.png\" alt=\"A skull has a pronounced sagittal crest, flaring cheekbones, and large hind teeth.\" width=\"381\" height=\"585\" \/><figcaption class=\"wp-caption-text\">Figure 9.12: Robust Australopithecines such as Paranthropus boisei had large molars and chewing muscles. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Paranthropus_boisei_skull.jpg\">Paranthropus boisei skull<\/a> by Durova is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/deed.en\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Between 3 mya and 1 mya, there seems to be differences in dietary strategy between different species of hominins designated as Australopithecines. A pattern of larger posterior dentition (even relative to the incisors and canines in the front of the mouth), thick enamel, and cranial evidence for extremely large chewing muscles is far more pronounced in a group known as the robust australopithecines. This pattern is extreme<span style=\"text-decoration: underline\">ly<\/span> relative to their earlier contemporaries or predecessors, the gracile australopithecines<strong>,<\/strong> and is certainly larger than those seen in early <em>Homo<\/em>, which emerged during this time. This pattern of incredibly large hind dentition (and very small anterior dentition) has led people to refer to robust australopithecines as <strong>megadont<\/strong> hominins (Figure 9.12).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Because of these differences, this section has been divided into \u201cgracile\u201d and \u201crobust\u201d Australopithecines, highlighting the morphological differences between the two groups (which many researchers have designated as separate genera: <em>Australopithecus<\/em> and <em>Paranthropus<\/em>, respectively) and then focusing on the individual species. It is worth noting, however, that not all researchers accept these clades as biologically or genetically distinct, with some researchers insisting that the relative gracile and robust features found in these species are due to parallel evolutionary events toward similar dietary niches.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Despite this genus\u2019 ancestral traits and small cranial capacity, all members show evidence of bipedal locomotion. It is generally accepted that <em>Australopithecus <\/em>species display varying degrees of arborealism along with bipedality.<\/span><\/p>\n<h3 class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Gracile Australopithecines<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">This section describes individual species from across Africa. These species are called \u201c<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1406\">gracile<\/a> <\/strong>australopithecines\u201d because of their smaller and less robust features compared to the divergent \u201c<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1408\"><strong>robust<\/strong><\/a>\u201d group. Numerous Australopithecine species have been named, but some are only based on a handful of fossil finds, whose designations are controversial.<\/span><\/p>\n<h4 class=\"import-Normal\"><em><span style=\"color: #000000\">East African Australopithecines<\/span><\/em><\/h4>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">East African Australopithecines are found throughout the EARS, and they include the earliest species associated with this genus. Numerous fossil-yielding sites, such as Olduvai, Turkana, and Laetoli, have excellent, datable stratigraphy, owing to the layers of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1362\"><strong>volcanic tufts<\/strong> <\/a> that have accumulated over millions of years. <span style=\"background-color: #ff9900\">These tufts may be dated using absolute dating techniques, such as Potassium-Argon dating (described in Chapter 7). This means that it is possible to know a relatively refined date for any fossil if the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1364\"><strong>context<\/strong> <\/a> (i.e., exact location) of that find is known.<\/span> Similarly, comparisons between the faunal assemblages of these stratigraphic layers have allowed researchers to chronologically identify environmental changes.<\/span><\/p>\n<figure style=\"width: 313px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image20-1-1.png\" alt=\"Occlusal view of an Au. anamensis mandible, with relatively large teeth, including canines.\" width=\"313\" height=\"313\" \/><figcaption class=\"wp-caption-text\">Figure 9.13: As seen in this mandible of KNM-KP 29281, Australopithecus anamensis had relatively large canine teeth. Credit: <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20anamensis\/KNM-KP%2029281\">Australopithecus anamensis: KNM-KP 29281 occlusal view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p><span style=\"color: #000000\">The earliest known Australopithecine is dated to 4.2 mya to 3.8 mya. <strong><em>Australopithecus anamensis<\/em><\/strong> (after \u201cAnam,\u201d meaning \u201clake\u201d from the Turkana region in Kenya; Leakey et al. 1995; Patterson and Howells 1967) is currently found from sites in the Turkana region (Kenya) and Middle Awash (Ethiopia; Figure 9.13). Recently, a 2019 find from Ethiopia, named MRD, after Miro Dora where it was found, was discovered by an Ethiopian herder named Ali Bereino. It is one of the most complete cranial finds of this species (Ward et al. 1999). A small brain size (370 cc), relatively large canines, projecting cheekbones, and earholes show more ancestral features as compared to those of more recent Australopithecines. The most important element discovered with this species is a fragment of a tibia (shinbone), which demonstrates features associated with weight transfer during bipedal walking. Similarly, the earliest found hominin femur belongs to this species. Ancestral traits in the upper limb (such as the humerus) indicate some retained arboreal locomotion.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Some researchers suggest that <em>Au. anamensis<\/em> is an intermediate form of the chronospecies that becomes <em>Au. afarensis<\/em>, evolving from <em>Ar. ramidus<\/em>. However, this is debated, with other researchers suggesting morphological similarities and affinities with more recent species instead. Almost 100 specimens, representing over 20 individuals, have been found to date (Leakey et al. 1995; McHenry 2009; Ward et al. 1999).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Au. afarensis<\/em><\/strong> is one of the oldest and most well-known australopithecine species and consists of a large number of fossil remains. <em>Au. afarensis<\/em> (which means \u201cfrom the Afar region\u201d) is dated to between 2.9 mya and 3.9 mya and is found in sites all along the EARS system, in Tanzania, Kenya, and Ethiopia (Figure 9.14). The most famous individual from this species is a partial female skeleton discovered in Hadar (Ethiopia), later nicknamed \u201cLucy,\u201d after the Beatles\u2019 song \u201cLucy in the Sky with Diamonds,\u201d which was played in celebration of the find (Johanson et al. 1978; Kimbel and Delezene 2009). This skeleton was found in 1974 by Donald Johanson and dates to approximately 3.2 mya. In addition, in 2002 a juvenile of the species was found by Zeresenay Alemseged and given the name \u201cSelam\u201d (meaning \u201cpeace,\u201d DIK 1-1), though it is popularly known as \u201cLucy\u2019s Child\u201d or as the \u201cDikika Child\u201d (Alemseged et al. 2006). Similarly, the \u201cLaetoli Footprints\u201d (discussed in Chapter 7; Hay and Leakey 1982; Leakey and Hay 1979) have drawn much attention.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 643px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-295 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.14.jpg\" alt=\"Two images of life-like reconstructions of female and male Au. afarensis.\" width=\"643\" height=\"322\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.14 a-b: Artistic reconstructions of Australopithecus afarensis by artist John Gurche. Female \u201cLucy\u201d is left and a male is on the right. Credit: a. <a class=\"rId106\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Australopithecus afarensis, \u201cLucy,\u201d adult female. Reconstruction based on AL-288-1 by artist John Gurche, front view close-up<\/a> by <a class=\"rId107\" href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?\u201d] is <a class=\"rId108\" href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>; b. <a class=\"rId109\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Australopithecus afarensis, adult male. Reconstruction based on <\/a><a class=\"rId110\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">AL444-2<\/a><a class=\"rId111\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\"> by John Gurche<\/a> by <a class=\"rId112\" href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?\u201d] is <a class=\"rId113\" href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>.<\/figcaption><\/figure>\n<figure style=\"width: 320px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image40.png\" alt=\"A partial skeleton includes bones of the cranium, mandible, and postcranium.\" width=\"320\" height=\"772\" \/><figcaption class=\"wp-caption-text\">Figure 9.15: The humanlike femoral neck, valgus knee, and bowl-shaped hip seen in the \u201cLucy\u201d skeleton indicates that Australopithecus afarensis was bipedal. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Lucy_blackbg.jpg\">Lucy blackbg<\/a> [AL 288-1, Australopithecus afarensis, cast from Museum national d'histoire naturelle, Paris] by 120 is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The canines and molars of <em>Au. afarensis<\/em> are reduced relative to great apes but are larger than those found in modern humans (indicative of a generalist diet); in addition, <em>Au. afarensis <\/em>has a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1366\"><strong>prognathic<\/strong> <\/a> face (the face below the eyes juts anteriorly) and robust facial features that indicate relatively strong chewing musculature (compared with <em>Homo<\/em>) but which are less extreme than in <em>Paranthropus<\/em>. Despite a reduction in canine size in this species, large overall size variation indicates high levels of sexual dimorphism.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Skeletal evidence indicates that this species was bipedal, as its pelvis and lower limb demonstrate a humanlike femoral neck, valgus knee, and bowl-shaped hip (Figure 9.15). <span style=\"background-color: #ccffcc\">More evidence of bipedalism is found in the footprints of this species<\/span>. <em>Au. afarensis<\/em> is associated with the Laetoli Footprints, <span style=\"background-color: #ff9900\">a 24-meter trackway of hominin fossil footprints preserved in volcanic ash discovered by Mary Leakey in Tanzania and dated to 3.5 mya to 3 mya. This set of prints is thought to have been produced by three bipedal individuals as there are no knuckle imprints, no opposable big toes, and a clear arch is present. The infants of this species are thought to have been more arboreal than the adults, as discovered through analyses of the foot bones of the Dikika Child dated to 3.32 mya (Alemseged et al. 2006).<\/span><\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Although not found in direct association with stone tools, potential evidence for cut marks on bones, found at Dikika, and dated to 3.39 mya indicates a possible temporal\/ geographic overlap between meat eating, tool use, and this species. However, this evidence is fiercely debated. Others have associated the cut marks with the earliest tool finds from Lomekwi, Kenya, temporally (3.3 mya) and in close geographic proximity to this species.<\/span><\/p>\n<h4 class=\"import-Normal\"><em><span style=\"color: #000000\">South African Australopithecines<\/span><\/em><\/h4>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Since the discovery of the Taung Child, there have been numerous Australopithecine discoveries from the region known as \u201cThe Cradle of Humankind,\u201d which was recently given UNESCO World Heritage Site status as \u201cThe Fossil Hominid Sites of South Africa.\u201d The limestone caves found in the Cradle allow for the excellent preservation of fossils. Past animals navigating the landscape and falling into cave openings, or caves used as dens by carnivores, led to the accumulation of deposits over millions of years. Many of the hominin fossils, encased in <strong>breccia<\/strong> (hard, calcareous sedimentary rock), are recently exposed from limestone quarries mined in the previous century. This means that extracting fossils requires excellent and detailed exposed work, often by a team of skilled technicians.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">While these sites have historically been difficult to date, with mixed assemblages accumulated over large time periods, advances in techniques such as uranium-series dating have allowed for greater accuracy. Historically, the excellent faunal record from East Africa has been used to compare sites based on <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1368\"><strong>relative dating<\/strong>,<\/a> whereby environmental and faunal changes and extinction events allow us to know which hominin finds are relatively younger or older than others.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The discovery of the Taung Child in 1924 (discussed in the Special Topic box \u201cThe Taung Child\u201d below) shifted the focus of palaeoanthropological research from Europe to Africa, although acceptance of this shift was slow (Broom 1947; Dart 1925). The species to which it is assigned, <strong><em>Australopithecus africanus<\/em><\/strong> (name meaning \u201cSouthern Ape of Africa\u201d), is currently dated to between 3.3 mya and 2.1 mya (Pickering and Kramers 2010), with discoveries from Sterkfontein, Taung, Makapansgat, and Gladysvale in South Africa (Figure 9.16). A relatively large brain (400 cc to 500 cc), small canines without an associated diastema, and more rounded cranium and smaller teeth than <em>Au. afarensis<\/em> indicate some derived traits. Similarly, the postcranial remains (in particular, the pelvis) indicate bipedalism. However, the sloping face and curved phalanges (indicative of retained arboreal locomotor abilities) show some ancestral features. Although not in direct association with stone tools, a 2015 study noted that the trabecular bone morphology of the hand was consistent with forceful tool manufacture and use, suggesting potential early tool abilities.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 570px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image19-1.png\" alt=\"A life-like reconstruction of the face of Australopithecus africanus, smiling in anterior view.\" width=\"570\" height=\"570\" \/><figcaption class=\"wp-caption-text\">Figure 9.16: An artistic reconstruction of Australopithecus africanus by John Gurche. Credit: <a href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Australopithecus africanus. Reconstruction based on STS 5 by John Gurche <\/a>by <a href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?] is <a href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Another famous <em>Au. africanus<\/em> skull (the skull of \u201cMrs. Ples\u201d) was previously attributed to <em>Plesianthropus transvaalensis<\/em><em>, <\/em>meaning \u201cnear human from the Transvaal,\u201d the old name for Gauteng Province, South Africa (Broom 1947, 1950). The name was shortened by contemporary journalists to \u201cPles\u201d (Figure 9.17). Due to the prevailing mores of the time, the assumed female found herself married, at least in name, and has become widely known as \u201cMrs. Ples.\u201d It was later reassigned to <em>Au. africanus<\/em> and is now argued by some to be a young male rather than an adult female cranium (Thackeray 2000, Thackeray et al. 2002).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 548px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-298 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.17.jpg\" alt=\"Four views of an ancient skull are shown on a black background.\" width=\"548\" height=\"548\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.17: The \u201cMrs. Ples\u201d brain case is small in size (like apes) but its face is less prognathic; its foramen magnum is positioned more like a modern human than an African apes. Credit: a. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 anterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 posterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 superior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; and d. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 lateral right view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In 2008, nine-year-old Matthew Berger, son of paleoanthropologist Lee Berger, noted a clavicle bone in some leftover mining breccia in the Malapa Fossil Site (South Africa). After rigorous studies, the species, <strong><em>Australopithecus sediba<\/em><\/strong> (meaning \u201cfountain\u201d or \u201cwellspring\u201d in the South African language of Sesotho), was named in 2010 (Figure 9.18; Berger et al. 2010). The first type specimen belongs to a juvenile male, Karabo (MH1), but the species is known from at least six partial skeletons, from infants through adults. These specimens are currently dated to 1.97 mya (Dirks et al. 2010). The discoverers have argued that <em>Au. sediba<\/em> shows mosaic features between <em>Au. africanus<\/em> and the genus, <em>Homo<\/em>, which potentially indicates a transitional species, although this is heavily debated. These features include a small brain size (<em>Australopithecus<\/em>-like; 420 cc to 450 cc) but gracile mandible and small teeth (<em>Homo<\/em>-like). Similarly, the postcranial skeletons are also said to have mosaic features: scientists have interpreted this mixture of traits (such as a robust ankle but evidence for an arch in the foot) as a transitional phase between a body previously adapted to arborealism (particularly in evidence from the bones of the wrist) to one that adapted to bipedal ground walking. Some researchers have argued that <em>Au. sediba<\/em> shows a modern hand morphology (shorter fingers and a longer thumb), indicating that adaptations to tool manufacture and use may be present in this species.<\/span><\/p>\n<figure style=\"width: 531px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image17-1.png\" alt=\"A beige-colored skull with no mandible on a black background has some missing teeth.\" width=\"531\" height=\"400\" \/><figcaption class=\"wp-caption-text\">Figure 9.18: Australopithecus sediba shows mosaic features between Au. africanus and Homo. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Australopithecus_sediba.JPG\">Australopithecus sediba<\/a>, photo by Brett Eloff courtesy <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Profberger\">Profberger<\/a> and <a href=\"https:\/\/en.wikipedia.org\/wiki\/University_of_the_Witwatersrand\">Wits University<\/a>, is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Another famous Australopithecine find from South Africa is that of the nearly complete skeleton now known as \u201cLittle Foot\u201d (Clarke 1998, 2013). Little Foot (StW 573) is potentially the earliest dated South African hominin fossil, dating to 3.7 mya, based on radiostopic techniques, although some argue that it is younger than 3 mya (Pickering and Kramers 2010). The name is jokingly in contrast to the cryptid species \u201cbigfoot\u201d and is named because the initial discovery of four ankle bones indicated bipedality. Little Foot was discovered by Ron Clarke in 1994, when he came across the ankle bones while sorting through monkey fossils in the University of Witwatersrand collections (Clarke and Tobias 1995). He asked Stephen Motsumi and Nkwane Molefe to identify the known records of the fossils, which allowed them to find the rest of the specimen within just days of searching the Sterkfontein Caves\u2019 Silberberg Grotto.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The discoverers of Little Foot insist that other fossil finds, previously identified as <em>Au. Africanus<\/em>, be placed in this new species based on shared ancestral traits with older East African Australopithecines (Clarke and Kuman 2019). These include features such as a relatively large brain size (408 cc), robust zygomatic arch, and a flatter midface. Furthermore, the discoverers have argued that the heavy anterior dental wear patterns, relatively large anterior dentition, and smaller hind dentition of this specimen more closely resemble that of <em>Au. anamensis<\/em> or <em>Au. afarensis<\/em>. It has thus been placed in the species <strong><em>Australopithecus prometheus<\/em><\/strong>. This species name refers to a previously defunct taxon named by Raymond Dart. The species designation was, through analyzing Little Foot, revived by Ron Clarke, who insists that many other fossil hominin specimens have prematurely been placed into <em>Au. africanus<\/em>. Others say that it is more likely that <em>Au. africanus<\/em> is a more variable species and not representative of two distinct species.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\"><em>Paranthropus<\/em> \u201cRobust\u201d Australopithecines<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In the robust australopithecines, the specialized nature of the teeth and masticatory system, such as flaring zygomatic arches (cheekbones), accommodate very large temporalis (chewing) muscles. These features also include a large, broad, dish-shaped face and and a large mandible with extremely large posterior dentition (referred to as megadonts) and hyper-thick enamel (Kimbel 2015; Lee-Thorp 2011; Wood 2010). Research has revolved around the shared adaptations of these \u201crobust\u201d australopithecines, linking their morphologies to a diet of hard and\/or tough foods (Brain 1967; Rak 1988). Some argued that the diet of the robust australopithecines was so specific that any change in environment would have accelerated their extinction. The generalist nature of the teeth of the gracile australopithecines, and of early <em>Homo<\/em>, would have made them more capable of adapting to environmental change. However, some have suggested that the features of the robust australopithecines might have developed as an effective response to what are known as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1370\"><strong>fallback <\/strong><strong>foods<\/strong><\/a> in hard times rather than indicating a lack of adaptability.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are currently three widely accepted robust australopithecus or, <em>Paranthropus<\/em>, species: <em>P. aethiopic<\/em><em>us<\/em>, which has more ancestral traits, and <em>P. boisei and P. robustus<\/em>, which are more derived in their features (Strait et al. 1997; Wood and Schroer 2017). These three species have been grouped together by a majority of scholars as a single genus as they share more derived features (are more closely related to each other; or, in other words, are <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1372\"><strong>monophyletic<\/strong><\/a>) than the other australopithecines (Grine 1988; Hlazo 2015; Strait et al. 1997; Wood 2010 ). While researchers have mostly agreed to use the umbrella term <em>Paranthropus<\/em>, there are those who disagree (Constantino and Wood 2004, 2007; Wood 2010).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">As a collective, this genus spans 2.7 mya to 1.0 mya, although the dates of the individual species differ. The earliest of the Paranthropus species, <strong><em>Paranthropus aethiopicus<\/em><\/strong>, is dated to between 2.7 mya and 2.3 mya and currently found in Tanzania, Kenya, and Ethiopia in the EARS system (Figure 9.19; Constantino and Wood 2007; Hlazo 2015; Kimbel 2015; Walker et al. 1986; White 1988). It is well known because of one specimen known as the \u201cBlack Skull\u201d (KNM\u2013WT 17000), so called because of the mineral manganese that stained it black during fossilization (Kimbel 2015). As with all robust Australopithecines, <em>P. aethiopicus<\/em> has the shared derived traits of large, flat premolars and molars; large, flaring zygomatic arches for accommodating large chewing muscles (the temporalis muscle); a sagittal crest (ridge on the top of the skull) for increased muscle attachment of the chewing muscles to the skull; and a robust mandible and supraorbital torus (brow ridge). However, only a few teeth have been found. A proximal tibia indicates bipedality and similar body size to <em>Au. afarensis<\/em>. In recent years, researchers have discovered and assigned a proximal tibia and juvenile cranium (L.338y-6) to the species (Wood and Boyle 2016).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 666px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-300 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.19.jpg\" alt=\"Five views of a beige partial skull on a black background.\" width=\"666\" height=\"444\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.19: The \u201cBlack Skull\u201d (Paranthropus aethiopicus) had a large sagittal crest and large, flared zygomatic arches that indicate it had large chewing muscles and a powerful biting force. Credit: a. <a class=\"rId156\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId157\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 anterior view<\/a> by \u00a9<a class=\"rId158\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId159\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId160\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId161\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId162\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 lateral right view<\/a> by \u00a9<a class=\"rId163\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId164\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId165\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a class=\"rId166\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId167\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 superior view<\/a> by \u00a9<a class=\"rId168\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId169\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId170\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; d. <a class=\"rId171\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId172\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 posterior view<\/a> by \u00a9<a class=\"rId173\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId174\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId175\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; e. <a class=\"rId176\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId177\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 inferior view<\/a> by \u00a9<a class=\"rId178\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId179\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId180\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">First attributed as <em>Zinjanthropus boisei<\/em> (with the first discovery going by the nickname \u201cZinj\u201d or sometimes \u201cNutcracker Man\u201d), <strong><em>Paranthropus boisei<\/em><\/strong> was discovered in 1959 by Mary Leakey (see Figure 9.20 and 9.21; Hay 1990; Leakey 1959). This \u201crobust\u201d australopith species is distributed across countries in East Africa at sites such as Kenya (Koobi Fora, West Turkana, and Chesowanja), Malawi (Malema-Chiwondo), Tanzania (Olduvai Gorge and Peninj), and Ethiopia (Omo River Basin and Konso). The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1374\">hypodigm<\/a><\/strong>, sample of fossils whose features define the group, has been found by researchers to date to roughly 2.4 mya to 1.4 mya. Due to the nature of its exaggerated, larger, and more robust features, <em>P. boisei <\/em>has been termed <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1376\"><strong>hyper-robust<\/strong><\/a>\u2014that is, even more heavily built than other robust species, with very large, flat posterior dentition (Kimbel 2015). Tools dated to 2.5 mya in Ethiopia have been argued to possibly belong to this species. Despite the cranial features of <em>P. boisei<\/em> indicating a tough diet of tubers, nuts, and seeds, isotopes indicate a diet high in C4 foods (e.g., grasses, such as sedges). Another famous specimen from this species is the Peninj mandible from Tanzania, found in 1964 by Kimoya Kimeu.<\/span><\/p>\n<figure style=\"width: 557px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image14-1.png\" alt=\"Life-like reconstruction of Paranthropus boisei.\" width=\"557\" height=\"557\" \/><figcaption class=\"wp-caption-text\">Figure 9.20: Artistic reconstruction of a Paranthropus boisei, male, by John Gurche. Credit: <a href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Paranthropus boisei, male. Reconstruction based on OH 5 and KNM-ER 406 by John Gurche<\/a> by <a href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?\u201d] is <a href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>.<\/figcaption><\/figure>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 565px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-302 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.21.jpg\" alt=\"Three views of an ancient skull are shown on a black background.\" width=\"565\" height=\"565\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.21: \u201cNutcracker Man\u201d (Paranthropus boisei) had hyper-robust features including very large dentition, flaring zygomatic arches, a broad concave face. It had a powerful and extremely efficient chewing force. Credit: <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20boisei\/OH%205\">Paranthropus boisei: OH 5 anterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20boisei\/OH%205\">Paranthropus boisei: OH 5 inferior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20boisei\/OH%205\">Paranthropus boisei: OH 5 posterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Paranthropus robustus<\/em><\/strong> was the first taxon to be discovered within the genus in Kromdraai B by a schoolboy named Gert Terblanche; subsequent fossil discoveries were made by researcher Robert Broom in 1938 (Figure 9.22; Broom 1938a, 1938b, 1950), with the holotype specimen TM 1517 (Broom 1938a, 1938b, 1950; Hlazo 2018). <em>Paranthropus robustus<\/em> dates approximately from 2.0 mya to 1 mya and is the only taxon from the genus to be discovered in South Africa. Several of these fossils are fragmentary in nature, distorted, and not well preserved because they have been recovered from quarry breccia using explosives. <em>P. robustus<\/em> features are neither as \u201chyper-robust\u201d as <em>P. boisei<\/em> nor as ancestral as <em>P. aethiopicus<\/em>; instead, they have been described as being less derived, more general features that are shared with both East African species (e.g., the sagittal crest and zygomatic flaring; Rak 1983; Walker and Leakey 1988). Enamel hypoplasia is also common in this species, possibly because of instability in the development of large, thick-enameled dentition.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 572px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-303 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.22.jpg\" alt=\"Four views of a beige-colored skull are shown on a black background.\" width=\"572\" height=\"619\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.22: SK 48, a Paranthropus robustus specimen, had less derived, more general features that were not as robust as P. boisei and not as ancestral as P. aethiopicus. Credit: a. <a class=\"rId208\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId209\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 anterior view<\/a> by \u00a9<a class=\"rId210\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId211\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId212\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId213\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId214\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 superior view<\/a> by \u00a9<a class=\"rId215\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId216\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId217\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a class=\"rId218\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId219\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 inferior view<\/a> by \u00a9<a class=\"rId220\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId221\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId222\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; d. <a class=\"rId223\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId224\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 lateral left view<\/a> by \u00a9<a class=\"rId225\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId226\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId227\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Comparisons between Gracile and Robust Australopiths<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Comparisons between gracile and robust australopithecines may indicate different phylogenetic groupings or parallel evolution in several species. In general, the robust australopithecines have large temporalis (chewing) muscles, as indicated by flaring zygomatic arches, sagittal crests, and robust mandibles (jawbones). Their hind dentition is large (megadont), with low cusps and thick enamel. Within the gracile australopithecines, researchers have debated the relatedness of the species, or even whether these species should be lumped together to represent more variable or polytypic species. Often researchers will attempt to draw chronospecific trajectories, with one taxon said to evolve into another over time.<\/span><\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Special Topic: The Taung Child<\/span><\/h2>\n<figure id=\"attachment_304\" aria-describedby=\"caption-attachment-304\" style=\"width: 570px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-304 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.23.jpg\" alt=\"An ancient skull in anterior and lateral views. One view shows an imprint of the brain.\" width=\"570\" height=\"285\" \/><figcaption id=\"caption-attachment-304\" class=\"wp-caption-text\">Figure 9.23: The Taung Child has a nearly complete face, mandible, and partial endocranial cast. Credit: a. <em>A<\/em><a class=\"rId230\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\"><em>ustralopithecus africanus<\/em><\/a><a class=\"rId231\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\">: Taung 1 anterior view<\/a> by \u00a9<a class=\"rId232\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId233\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId234\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId235\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\"><em>australopithecus africanus<\/em><\/a><a class=\"rId236\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\">: Taung 1 lateral right view<\/a> by \u00a9<a class=\"rId237\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId238\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId239\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">The well-known fossil of a juvenile <em>Australopithecine<\/em>, the \u201cTaung Child,\u201d was the first early hominin evidence ever discovered and was the first to demonstrate our common human heritage in Africa (Figure 9.23; Dart 1925). The tiny facial skeleton and natural endocast were discovered in 1924 by a local quarryman in the North West Province in South Africa and were painstakingly removed from the surrounding cement-like breccia by Raymond Dart using his wife\u2019s knitting needles. When first shared with the scientific community in 1925, it was discounted as being nothing more than a young monkey of some kind. Prevailing biases of the time made it too difficult to contemplate that this small-brained hominin could have anything to do with our own history. The fact that it was discovered in Africa simply served to strengthen this bias.<\/span><\/p>\n<\/div>\n<p>&nbsp;<\/p>\n<h2><span style=\"color: #000000\">Early Tool Use and Technology<br \/>\n<\/span><\/h2>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Early Stone Age Technology (ESA)<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1378\">Early Stone Age (ESA)<\/a><\/strong> marks the beginning of recognizable technology made by our human ancestors. Stone-tool (or <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1380\"><strong>lithic<\/strong><\/a>) technology is defined by the fracturing of rocks and the manufacture of tools through a process called <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1382\"> <strong>knapping<\/strong><\/a>. The Stone Age lasted for more than 3 million years and is broken up into chronological periods called the Early (ESA), Middle (MSA), and Later Stone Ages (LSA). Each period is further broken up into a different <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1384\">techno-complex<\/a><\/strong>, a term encompassing multiple <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1386\">assemblages<\/a><\/strong> (collections of artifacts) that share similar traits in terms of artifact production and morphology. The ESA spanned the largest technological time period of human innovation from over 3 million years ago to around 300,000 years ago and is associated almost entirely with hominin species prior to modern <em>Homo sapiens. <\/em>As the ESA advanced, stone tool makers (known as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1390\"><strong>knappers<\/strong><\/a>) began to change the ways they detached <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1388\">flakes<\/a><\/strong> and eventually were able to shape artifacts into functional tools. These advances in technology go together with the developments in human evolution and cognition, dispersal of populations across the African continent and the world, and climatic changes.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In order to understand the ESA, it is important to consider that not all assemblages are exactly the same within each techno-complex: one can have multiple phases and traditions at different sites (Lombard et al. 2012). However, there is an overarching commonality between them. Within stone tool assemblages, both flakes or <strong>cores<\/strong> (the rocks from which flakes are removed) are used as tools.<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1392\"> <strong>Large Cutting Tools (LCTs)<\/strong><\/a> are tools that are shaped to have functional edges. It is important to note that the information presented here is a small fraction of what is known about the ESA, and there are ongoing debates and discoveries within archaeology.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Currently, the oldest-known stone tools, which form the techno-complex the Lomekwian, date to 3.3 mya (Harmand et al. 2015; Toth 1985). They were found at a site called Lomekwi 3 in Kenya. This techno-complex is the most recently defined and pushed back the oldest-known date for lithic technology. There is only one known site thus far and, due to the age of the site, it is associated with species prior to <em>Homo<\/em>, such as <em>Kenyanthropus platyops.<\/em> Flakes were produced through indirect percussion, whereby the knappers held a rock and hit it against another rock resting on the ground. The pieces are very chunky and do not display the same fracture patterns seen in later techno-complexes. Lomekwian knappers likely aimed to get a sharp-edged piece on a flake, which would have been functional, although the specific function is currently unknown.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Stone tool use, however, is not only understood through the direct discovery of the tools. Cut marks on fossilized animal bones may illuminate the functionality of stone tools. In one controversial study in 2010, researchers argued that cut marks on a pair of animal bones from Dikika (Ethiopia), dated to 3.4 mya, were from stone tools. The discoverers suggested that they be more securely associated, temporally, with <em>Au. afarensis<\/em>. However, others have noted that these marks are consistent with teeth marks from crocodiles and other carnivores.<\/span><\/p>\n<figure style=\"width: 324px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image29-1.png\" alt=\"A technical line drawing of an Oldowan chopper.\" width=\"324\" height=\"275\" \/><figcaption class=\"wp-caption-text\">Figure 9.24: Some scholars believe that some genera explored in this chapter were capable of producing more complex stone tools (Oldowan). Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Olduwan_Industry_Chopper_2.jpeg\">Olduwan Industry Chopper 2<\/a> by Emmyanne29 is under a <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/legalcode\">CC0 1.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The Oldowan techno-complex is far more established in the scientific literature (Leakey 1971). It is called the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1394\">Oldowan<\/a><\/strong> because it was originally discovered in Olduvai Gorge, Tanzania, but the oldest assemblage is from Gona in Ethiopia, dated to 2.6 mya (Semaw 2000). The techno-complex is defined as a core and flake industry. Like the Lomekwian, there was an aim to get sharp-edged flakes, but this was achieved through a different production method. Knappers were able to actively hold or manipulate the core being knapped, which they could directly hit using a hammerstone. This technique is known as free-hand percussion, and it demonstrates an understanding of fracture mechanics. It has long been argued that the Oldowan hominins were skillful in tool manufacture.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Because Oldowan knapping requires skill, earlier researchers have attributed these tools to members of our genus, <em>Homo<\/em>. However, some have argued that these tools are in more direct association with hominins in the genera described in this chapter (Figure 9.24).<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Invisible Tool Manufacture and Use<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The vast majority of our understanding of these early hominins comes from fossils and reconstructed paleoenvironments. It is only from 3 mya when we can start \u201clooking into their minds\u201d and lifestyles by analyzing their manufacture and use of stone tools. However, the vast majority of tool use in primates (and, one can argue, in humans) is not with durable materials like stone. All of our extant great ape relatives have been observed using sticks, leaves, and other materials for some secondary purpose (to wade across rivers, to \u201cfish\u201d for termites, or to absorb water for drinking). It is possible that the majority of early hominin tool use and manufacture may be invisible to us because of this preservation bias.<\/span><\/p>\n<h2 class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000;background-color: #ff00ff\">Chapter Summary<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000;background-color: #ff00ff\">The fossil record of our earliest hominin relatives has allowed paleoanthropologists to unpack some of the mysteries of our evolution. We now know that traits associated with bipedalism evolved before other \u201chuman-like\u201d traits, even though the first hominins were still very capable of arboreal locomotion. We also know that, for much of this time, hominin taxa were diverse in the way they looked and what they ate, and they were widely distributed across the African continent. And we know that the environments in which these hominins lived underwent many changes over this time during several warming and cooling phases.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;background-color: #ff00ff\">Yet this knowledge has opened up many new mysteries. We still need to better differentiate some taxa. In addition, there are ongoing debates about why certain traits evolved and what they meant for the extinction of some of our relatives (like the robust australopiths). The capabilities of these early hominins with respect to tool use and manufacture is also still uncertain.<\/span><\/p>\n<h2 class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Hominin Species Summaries<br \/>\n<\/span><\/h2>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Sahelanthropus tchadensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">7 mya to 6 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Chad<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The initial discovery, made in 2001.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">360 cc average<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Smaller than in extant great apes; larger and pointier than in humans. Canines worn at the tips.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A short cranial base and a foramen magnum (hole in which the spinal cord enters the cranium) that is more humanlike in positioning; has been argued to indicate upright walking.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Currently little published postcranial material.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The extent to which this hominin was bipedal is currently heavily debated. If so, it would indicate an arboreal bipedal ancestor of hominins, not a knuckle-walker like chimpanzees.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Orrorin tugenensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">6 mya to 5.7 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Tugen Hills (Kenya)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Original discovery in 2000.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Smaller cheek teeth (molars and premolars) than even more recent hominins (i.e., derived), thick enamel, and reduced, but apelike, canines.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Not many found<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Fragmentary leg, arm, and finger bones have been found. Indicates bipedal locomotion.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Potential toolmaking capability based on hand morphology, but nothing found directly.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">This is the earliest species that clearly indicates adaptations for bipedal locomotion.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Ardipithecus kadabba<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">5.2 mya to 5.8 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Middle Awash (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Discovered by Yohannes Haile-Selassie in 1997.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Larger hind dentition than in modern chimpanzees. Thick enamel and larger canines than in later hominins.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A large hallux (big toe) bone indicates a bipedal \u201cpush off.\u201d<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Faunal evidence indicates a mixed grassland\/woodland environment.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Ardipithecus ramidus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">4.4 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Middle Awash region and Gona (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">A partial female skeleton nicknamed \u201cArdi\u201d (ARA-VP-6\/500) (found in 1994).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">300 cc to 350 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Little differences between the canines of males and females (small sexual dimorphism).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Midfacial projection, slightly prognathic. Cheekbones less flared and robust than in later hominins.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Ardi demonstrates a mosaic of ancestral and derived characteristics in the postcrania. For instance, an opposable big toe similar to chimpanzees (i.e., more ancestral), which could have aided in climbing trees effectively. However, the pelvis and hip show that she could walk upright (i.e., it is derived), supporting her hominin status.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">None directly associated<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">Over 110 specimens from Aramis<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus anamensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">4.2 mya to 3.8 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Turkana region (Kenya); Middle Awash (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A 2019 find from Ethiopia, named MRD.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">370 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Relatively large canines compared with more recent Australopithecines.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Projecting cheekbones and ancestral earholes.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lower limb bones (tibia and femur) indicate bipedality; arboreal features in upper limb bones (humerus) found.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Almost 100 specimens, representing over 20 individuals, have been found to date.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus afarensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">3.9 mya to 2.9 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Afar Region, Omo, Maka, Fejej, and Belohdelie (Ethiopia); Laetoli (Tanzania); Koobi Fora (Kenya)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lucy (discovery: 1974), Selam (Dikika Child, discovery: 2000), Laetoli Footprints (discovery: 1976).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">380 cc to 430 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Reduced canines and molars relative to great apes but larger than in modern humans.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Prognathic face, facial features indicate relatively strong chewing musculature (compared with <em>Homo<\/em>) but less extreme than in <em>Paranthropus<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clear evidence for bipedalism from lower limb postcranial bones. Laetoli Footprints indicate humanlike walking. Dikika Child bones indicate retained ancestral arboreal traits in the postcrania.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">None directly, but close in age and proximity to controversial cut marks at Dikika and early tools in Lomekwi.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Au. afarensis<\/em> is one of the oldest and most well-known australopithecine species and consists of a large number of fossil remains.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus bahrelghazali<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">3.6 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Chad<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">\u201cAbel,\u201d the holotype (discovery: 1995).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Arguably within range of variation of <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus prometheus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">3.7 mya (debated)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">Sterkfontein (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">\u201cLittle Foot\u201d (StW 573) (discovery: 1994)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">408 cc (Little Foot estimate)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">Heavy anterior dental wear patterns, relatively large anterior dentition and smaller hind dentition, similar to <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">Relatively larger brain size, robust zygomatic arch, and a flatter midface.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">The initial discovery of four ankle bones indicated bipedality.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">Highly debated new species designation.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus <\/em><em>deyiremada<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">3.5 mya to 3.3 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Woranso-Mille (Afar region, Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">First fossil mandible bones were discovered in 2011 in the Afar region of Ethiopia by Yohannes Haile-Selassie.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Smaller teeth with thicker enamel than seen in <em>Au. afarensis<\/em>, with a potentially hardier diet.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Larger mandible and more projecting cheekbones than in <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Contested species designation; arguably a member of <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Kenyanthopus<\/em><em> platyops<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">3.5 mya to 3.2 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Lake Turkana (Kenya)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">KNM\u2013WT 40000 (discovered 1999)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Difficult to determine but appears within the range of <em>Australopithecus afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Small molars\/dentition (<em>Homo<\/em>-like characteristic)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Flatter (i.e., orthognathic) face<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Some have associated the earliest tool finds from Lomekwi, Kenya, temporally (3.3 mya) and in close geographic proximity to this species\/specimen.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Taxonomic placing of this species is quite divided. The discoverers have argued that this species is ancestral to <em>Homo<\/em>, in particular to <em>Homo <\/em><em>ruldolfensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Australopithecus africanus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">3.3 mya to 2.1 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Sterkfontein, Taung, Makapansgat, Gladysvale (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Taung Child (discovery in 1994), \u201cMrs. Ples\u201d (discover in 1947), Little Foot (arguable; discovery in 1994).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">400 cc to 500 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Smaller teeth (derived) relative to <em>Au. afarensis<\/em>. Small canines with no diastema.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">A rounder skull compared with <em>Au. afarensis<\/em> in East Africa. A sloping face (ancestral).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Similar postcranial evidence for bipedal locomotion (derived pelvis) with retained arboreal locomotion, e.g., curved phalanges (fingers), as seen in <em>Au. afarensis.<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">None with direct evidence.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">A 2015 study noted that the trabecular bone morphology of the hand was consistent with forceful tool manufacture and use, suggesting potential early tool abilities.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus garhi<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.5 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Middle Awash (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">450 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Larger hind dentition than seen in other gracile Australopithecines.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A femur of a fragmentary partial skeleton, argued to belong to <em>Au. garhi<\/em>, indicates this species may be longer-limbed than <em>Au. afarensis<\/em>, although still able to move arboreally.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Crude stone tools resembling Oldowan (described later) have been found in association with <em>Au. garhi<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">This species is not well documented or understood and is based on only a few fossil specimens.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Paranthropus aethiopicus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.7 mya to 2.3 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">West Turkana (Kenya); Laetoli (Tanzania); Omo River Basin (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The \u201cBlack Skull\u201d (KNM\u2013WT 17000) (discovery 1985).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain Size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">410 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>P. aethiopicus<\/em> has the shared derived traits of large flat premolars and molars, although few teeth have been found.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Large flaring zygomatic arches for accommodating large chewing muscles (the temporalis muscle), a sagittal crest for increased muscle attachment of the chewing muscles to the skull, and a robust mandible and supraorbital torus (brow ridge).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A proximal tibia indicates bipedality and similar size to <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The \u201cBlack Skull\u201d is so called because of the mineral manganese that stained it black during fossilization.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Paranthropus boisei<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.4 mya to 1.4 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Koobi Fora, West Turkana, and Chesowanja (Kenya); Malema-Chiwondo (Malawi), Olduvai Gorge and Peninj (Tanzania); and Omo River basin and Konso (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">\u201cZinj,\u201d or sometimes \u201cNutcracker Man\u201d (OH5), in 1959 by Mary Leakey. The Peninj mandible from Tanzania, found in 1964 by Kimoya Kimeu.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">500 cc to 550 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Very large, flat posterior dentition (largest of all hominins currently known). Much smaller anterior dentition. Very thick dental enamel.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Indications of very large chewing muscles (e.g., flaring zygomatic arches and a large sagittal crest).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Evidence for high variability and sexual dimorphism, with estimates of males at 1.37 meters tall and females at 1.24 meters.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Richard Leakey and Bernard Wood have both suggested that<em> P. boisei<\/em> could have made and used stone tools. Tools dated to 2.5 mya in Ethiopia have been argued to possibly belong to this species.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Despite the cranial features of <em>P. boisei<\/em> indicating a tough diet of tubers, nuts, and seeds, isotopes indicate a diet high in C4 foods (e.g., grasses, such as sedges). This differs from what is seen in<em> P. robustus<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Australopithecus sediba<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">1.97 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Malapa Fossil Site (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Karabo (MH1) (discovery in 2008)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">420 cc to 450 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Small dentition with Australopithecine cusp-spacing.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Small brain size (<em>Australopithecus<\/em>-like) but gracile mandible (<em>Homo<\/em>-like).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Scientists have interpreted this mixture of traits (such as a robust ankle but evidence for an arch in the foot) as a transitional phase between a body previously adapted to arborealism (tree climbing, particularly in evidence from the bones of the wrist) to one that adapted to bipedal ground walking.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">None of direct association, but some have argued that a modern hand morphology (shorter fingers and a longer thumb) means that adaptations to tool manufacture and use may be present in this species.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">It was first discovered through a clavicle bone in 2008 by nine-year-old Matthew Berger, son of paleoanthropologist Lee Berger.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Paranthropus robustus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.3 mya to 1 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kromdraai B, Swartkrans, Gondolin, Drimolen, and Coopers Cave (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">SK48 (original skull)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">410 cc to 530 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Large posterior teeth with thick enamel, consistent with other Robust Australopithecines. Enamel hypoplasia is also common in this species, possibly because of instability in the development of large, thick enameled dentition.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>P. robustus<\/em> features are neither as \u201chyper-robust\u201d as <em>P. boisei<\/em> or as ancestral in features as <em>P. aethiopicus<\/em>. They have been described as less derived, more general features that are shared with both East African species (e.g., the sagittal crest and zygomatic flaring).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Reconstructions indicate sexual dimorphism.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Several of these fossils are fragmentary in nature, distorted, and not well preserved, because they have been recovered from quarry breccia using explosives.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\"><strong><span style=\"color: #000000\">Review Questions<br \/>\n<\/span><\/strong><\/h2>\n<ul>\n<li class=\"import-Normal\"><span style=\"color: #000000\">What is the difference between a \u201cderived\u201d versus an \u201cancestral\u201d trait? Give an example of both, seen in <em>Au. afarensis<\/em>.<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">Which of the paleoenvironment hypotheses have been used to describe early hominin diversity, and which have been used to describe bipedalism?<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">Which anatomical features for bipedalism do we see in early hominins?<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">Describe the dentition of gracile and robust australopithecines. What might these tell us about their diets?<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">List the hominin species argued to be associated with stone tool technologies. Are you convinced of these associations? Why\/why not?<\/span><\/li>\n<\/ul>\n<\/div>\n<h2><span style=\"color: #000000\">Key Terms<\/span><\/h2>\n<p><span style=\"color: #000000\"><strong>Arboreal:<\/strong> Related to trees or woodland.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Aridification:<\/strong> Becoming increasingly arid or dry, as related to the climate or environment.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Aridity Hypothesis:<\/strong> The hypothesis that long-term aridification and expansion of savannah biomes were drivers in diversification in early hominin evolution.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Assemblage:<\/strong> A collection demonstrating a pattern. Often pertaining to a site or region.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Bipedalism:<\/strong> The locomotor ability to walk on two legs.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Breccia:<\/strong> Hard, calcareous sedimentary rock.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Canines:<\/strong> The pointy teeth just next to the incisors, in the front of the mouth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cheek teeth:<\/strong> Or hind dentition (molars and premolars).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Chronospecies:<\/strong> Species that are said to evolve into another species, in a linear fashion, over time.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Clade:<\/strong> A group of species or taxa with a shared common ancestor.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cladistics:<\/strong> The field of grouping organisms into those with shared ancestry.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Context:<\/strong> As pertaining to palaeoanthropology, this term refers to the place where an artifact or fossil is found.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cores:<\/strong> The remains of a rock that has been flaked or knapped.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cusps:<\/strong> The ridges or \u201cbumps\u201d on the teeth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Dental formula:<\/strong> A technique to describe the number of incisors, canines, premolars, and molars in each quadrant of the mouth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Derived traits:<\/strong> Newly evolved traits that differ from those seen in the ancestor.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Diastema:<\/strong> A tooth gap between the incisors and canines.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Early Stone Age (ESA):<\/strong> The earliest-described archaeological period in which we start seeing stone-tool technology.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>East African Rift System (EARS):<\/strong> This term is often used to refer to the Rift Valley, expanding from Malawi to Ethiopia. This active geological structure is responsible for much of the visibility of the paleoanthropological record in East Africa.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Enamel:<\/strong> The highly mineralized outer layer of the tooth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Encephalization:<\/strong> Expansion of the brain.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Extant:<\/strong> Currently living\u2014i.e., not extinct.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Fallback foods:<\/strong> Foods that may not be preferred by an animal (e.g., foods that are not nutritionally dense) but that are essential for survival in times of stress or scarcity.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Fauna:<\/strong> The animals of a particular region, habitat, or geological period.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Faunal assemblages:<\/strong> Collections of fossils of the animals found at a site.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Faunal turnover:<\/strong> The rate at which species go extinct and are replaced with new species.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Flake:<\/strong> The piece knocked off of a stone core during the manufacture of a tool, which may be used as a stone tool.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Flora:<\/strong> The plants of a particular region, habitat, or geological period.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Folivorous:<\/strong> Foliage-eating.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Foramen magnum:<\/strong> The large hole (foramen) at the base of the cranium, through which the spinal cord enters the skull.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Fossil:<\/strong> The remains or impression of an organism from the past.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Frugivorous:<\/strong> Fruit-eating.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Generalist:<\/strong> A species that can thrive in a wide variety of habitats and can have a varied diet.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Glacial:<\/strong> Colder, drier periods during an ice age when there is more ice trapped at the poles.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Gracile:<\/strong> Slender, less rugged, or pronounced features.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hallux:<\/strong> The big toe.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Holotype:<\/strong> A single specimen from which a species or taxon is described or named.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hominin:<\/strong> A primate category that includes humans and our fossil relatives since our divergence from extant great apes.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Honing P3:<\/strong> The mandibular premolar alongside the canine (in primates, the P3), which is angled to give space for (and sharpen) the upper canines.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hyper-robust:<\/strong> Even more robust than considered normal in the Paranthropus genus.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hypodigm:<\/strong> A sample (here, fossil) from which researchers extrapolate features of a population.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Incisiform:<\/strong> An adjective referring to a canine that appears more incisor-like in morphology.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Incisors:<\/strong> The teeth in the front of the mouth, used to bite off food.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Interglacial:<\/strong> A period of milder climate in between two glacial periods.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Isotopes:<\/strong> Two or more forms of the same element that contain equal numbers of protons but different numbers of neutrons, giving them the same chemical properties but different atomic masses.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Knappers:<\/strong> The people who fractured rocks in order to manufacture tools.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Knapping:<\/strong> The fracturing of rocks for the manufacture of tools.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Large Cutting Tool (LCT):<\/strong> A tool that is shaped to have functional edges.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Last Common Ancestor (LCA):<\/strong> The hypothetical final ancestor (or ancestral population) of two or more taxa before their divergence.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Lithic:<\/strong> Relating to stone (here to stone tools).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Lumbar lordosis:<\/strong> The inward curving of the lower (lumbar) parts of the spine. The lower curve in the human S-shaped spine.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Lumpers:<\/strong> Researchers who prefer to lump variable specimens into a single species or taxon and who feel high levels of variation is biologically real.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Megadont:<\/strong> An organism with extremely large dentition compared with body size.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Metacarpals:<\/strong> The long bones of the hand that connect to the phalanges (finger bones).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Molars:<\/strong> The largest, most posterior of the hind dentition.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Monophyletic:<\/strong> A taxon or group of taxa descended from a common ancestor that is not shared with another taxon or group.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Morphology:<\/strong> The study of the form or size and shape of things; in this case, skeletal parts.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Mosaic evolution:<\/strong> The concept that evolutionary change does not occur homogeneously throughout the body in organisms.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Obligate bipedalism:<\/strong> Where the primary form of locomotion for an organism is bipedal.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Occlude:<\/strong> When the teeth from the maxilla come into contact with the teeth in the mandible.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Oldowan:<\/strong> Lower Paleolithic, the earliest stone tool culture.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Orthognathic:<\/strong> The face below the eyes is relatively flat and does not jut out anteriorly.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Paleoanthropologists:<\/strong> Researchers that study human evolution.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Paleoenvironment:<\/strong> An environment from a period in the Earth\u2019s geological past.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Parabolic:<\/strong> Like a parabola (parabola-shaped).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Phalanges:<\/strong> Long bones in the hand and fingers.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Phylogenetics:<\/strong> The study of phylogeny.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Phylogeny:<\/strong> The study of the evolutionary relationships between groups of organisms.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Pliocene:<\/strong> A geological epoch between the Miocene and Pleistocene.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Polytypic:<\/strong> In reference to taxonomy, having two or more group variants capable of interacting and breeding biologically but having morphological population differences.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Postcranium:<\/strong> The skeleton below the cranium (head).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Premolars:<\/strong> The smallest of the hind teeth, behind the canines.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Procumbent:<\/strong> In reference to incisors, tilting forward.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Prognathic:<\/strong> In reference to the face, the area below the eyes juts anteriorly.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Quaternary Ice Age:<\/strong> The most recent geological time period, which includes the Pleistocene and Holocene Epochs and which is defined by the cyclicity of increasing and decreasing ice sheets at the poles.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Relative dating:<\/strong> Dating techniques that refer to a temporal sequence (i.e., older or younger than others in the reference) and do not estimate actual or absolute dates.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Robust:<\/strong> Rugged or exaggerated features.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Site:<\/strong> A place in which evidence of past societies\/species\/activities may be observed through archaeological or paleontological practice.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Specialist:<\/strong> A specialist species can thrive only in a narrow range of environmental conditions or has a limited diet.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Splitters:<\/strong> Researchers who prefer to split a highly variable taxon into multiple groups or species.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Taxa:<\/strong> Plural of taxon, a taxonomic group such as species, genus, or family.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Taxonomy:<\/strong> The science of grouping and classifying organisms.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Techno-complex:<\/strong> A term encompassing multiple assemblages that share similar traits in terms of artifact production and morphology.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Thermoregulation:<\/strong> Maintaining body temperature through physiologically cooling or warming the body.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Ungulates:<\/strong> Hoofed mammals\u2014e.g., cows and kudu.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Volcanic tufts:<\/strong> Rock made from ash from volcanic eruptions in the past.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Valgus knee:<\/strong> The angle of the knee between the femur and tibia, which allows for weight distribution to be angled closer to the point above the center of gravity (i.e., between the feet) in bipeds.<\/span><\/p>\n<h2 class=\"import-Normal\"><strong><span style=\"color: #000000\">About the Authors<br \/>\n<\/span><\/strong><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image31.jpg\" alt=\"A woman with short blonde hair smiles at the camera.\" width=\"311\" height=\"311\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Kerryn Warren, Ph.D.<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Grad Coach International, <a class=\"rId245\" style=\"color: #000000\" href=\"mailto:kerryn.warren@gmail.com\">kerryn.warren@gmail.com<\/a><\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kerryn Warren is a dissertation coach at Grad Coach International and is passionate about stimulating research thinking in students of all levels. She has lectured on multiple topics, including archaeology and human evolution, with her research and science communication interests including hybridization in the hominin fossil record (stemming from research from her Ph.D.) and understanding how evolution is taught in South African schools. She also worked as one of the \u201cUnderground Astronauts,\u201d selected to excavate <em>Homo naledi <\/em>remains from the Rising Star Cave System in the Cradle of Humankind.<\/span><\/p>\n<p>&nbsp;<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image11-2.jpg\" alt=\"A woman with short brown hair smiles at the camera.\" width=\"312\" height=\"306\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">K. Lindsay Hunter, M.A., Ph.D. candidate<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">CARTA, k.lindsay.hunter@gmail.com<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lindsay Hunter is a trained palaeoanthropologist who uses her more than 15 years of experience to make sense of the distant past of our species to build a better future. She received her master\u2019s degree in biological anthropology from the University of Iowa and is completing her Ph.D. in archaeology at the University of the Witwatersrand in Johannesburg, South Africa. She has studied fossil and human bone collections across five continents with major grant support from the National Science Foundation (United States) and the Wenner-Gren Foundation for Anthropological Research. As a National Geographic Explorer, Lindsay developed and managed the National Geographic\u2013sponsored Umsuka Public Palaeoanthropology Project in the Cradle of Humankind World Heritage Site (CoH WHS) in South Africa from within Westbury Township, Johannesburg, between 2016\u20132019. She currently serves as the Community Engagement &amp; Advancement Director for CARTA: The UC San Diego\/Salk Institute Center for Academic Research and Training in Anthropogeny in La Jolla, California.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image25-2.jpg\" alt=\"A woman with black hair stands in a hole in the ground.\" width=\"295\" height=\"339\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Navashni Naidoo, M.Sc.<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">University of Cape Town, nnaidoo2@illinois.edu<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Navashni Naidoo is a researcher at Nelson Mandela University, lecturing on physical geology. She completed her Master\u2019s in Science in Archaeology in 2017 at the University of Cape Town. Her research interests include developing paleoenvironmental proxies suited to the African continent, behavioral ecology, and engaging with community-driven archaeological projects. She has excavated at Stone Age sites across Southern Africa and East Africa. Navashni is currently pursuing a PhD in the Department of Anthropology at the University of Illinois.<\/span><\/p>\n<p>&nbsp;<\/p>\n<p>&nbsp;<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image47.jpg\" alt=\"A man with black hair and dark brown eyes looks at the camera. \" width=\"294\" height=\"294\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Silindokuhle Mavuso, M.Sc.<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">University of Witwatersrand, S.muvaso@ru.ac.za<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Silindokuhle has always been curious about the world around him and how it has been shaped. He is a lecturer at Rhodes University of Witwatersrand (Wits), and conducts research on palaeoenvironmental reconstruction and change of the northeastern Turkana Basin\u2019s Pleistocene sequence. Silindokuhle began his education with a B.Sc. (Geology, Archaeology, and Environmental and Geographical Sciences) from the University of Cape Town before moving to Wits for a B.Sc. Honors (geology and paleontology) and M.Sc. in geology. He is currently concluding his PhD Studies. During this time, he has gained more training as a Koobi Fora Fieldschool fellow (Kenya) as well as an Erasmus Mundus scholar (France). Silindokuhle is a Plio-Pleistocene geologist with a specific focus on identifying and explaining past environments that are associated with early human life and development through time. He is interested in a wide range of disciplines such as micromorphology, sedimentology, geochemistry, geochronology, and sequence stratigraphy. He has worked with teams from significant eastern and southern African hominid sites including Elandsfontein, Rising Star, Sterkfontein, Gondolin, Laetoli, Olduvai, and Koobi Fora.<\/span><\/p>\n<h2 class=\"import-Normal\"><strong><span style=\"color: #000000\">For Further Exploration<br \/>\n<\/span><\/strong><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/humanorigins.si.edu\/evidence\">The Smithsonian Institution website<\/a> hosts descriptions of fossil species, an interactive timeline, and much more.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.maropeng.co.za\/content\/page\/human-evolution\">The Maropeng Museum website<\/a> hosts a wealth of information regarding South African Fossil Bearing sites in the Cradle of Humankind<strong>.<\/strong><\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/perot-museum.imgix.net\/2019-08-naledi-sediba-quick-comparison.pdf\">This quick comparison between <em>Homo naledi<\/em> and <em>Australopithecus sediba<\/em><\/a> from the Perot Museum.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.dropbox.com\/s\/l1d2hv42psj21y9\/Braided%20Stream-1920.mp4?dl=0\">This explanation of the braided stream<\/a> by the Perot Museum.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.hetmp.com\/\">A collation of 3-D files for visualizing<\/a> (or even 3-D printing) for homes, schools, and universities.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.pbslearningmedia.org\/resource\/tdc02.sci.life.evo.lp_humanevo\/human-evolution.\">PBS learning materials<\/a>, including videos and diagrams of the Laetoli footprints, bipedalism, and fossils.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A wealth of <a href=\"https:\/\/australianmuseum.net.au\/learn\/science\/human-evolution\/\">information from the Australian Museum website<\/a>, including species descriptions, family trees, and explanations of bipedalism and diet<strong>.<\/strong><\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\"><strong>References<\/strong><\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Alemseged, Zeresenay, Fred Spoor, William H. Kimbel, Ren\u00e9 Bobe, Denis Geraads, Denn\u00e9 Reed, and Jonathan G. Wynn. 2006. \u201cA Juvenile Early Hominin Skeleton from Dikika, Ethiopia.\u201d <em>Nature<\/em> 443 (7109): 296\u2013301.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Asfaw, Berhane, Tim White, Owen Lovejoy, Bruce Latimer, Scott Simpson, and Gen Suwa. 1999. \u201c<em>Australopithecus garhi<\/em>: A New Species of Early Hominid from Ethiopia.\u201d <em>Science<\/em> 284 (5414): 629\u2013635.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Behrensmeyer, Anna K., Nancy E. Todd, Richard Potts, and Geraldine E. McBrinn. 1997. \u201cLate Pliocene Faunal Turnover in the Turkana Basin, Kenya, and Ethiopia.\u201d <em>Science<\/em> 278 (5343): 637\u2013640.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Berger, Lee R., Darryl J. De Ruiter, Steven E. Churchill, Peter Schmid, Kristian J. Carlson, Paul HGM Dirks, and Job M. Kibii. 2010. \u201c<em>Australopithecus sediba<\/em>: A New Species of <em>Homo<\/em>-like Australopith from South Africa.\u201d <em>Science<\/em> 328 (5975): 195\u2013204.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Bobe, Ren\u00e9, and Anna K. Behrensmeyer. 2004. \u201cThe Expansion of Grassland Ecosystems in Africa in Relation to Mammalian Evolution and the Origin of the Genus <em>Homo<\/em>.\u201d <em>Palaeogeography, Palaeoclimatology, Palaeoecology<\/em> 207 (3\u20134): 399\u2013420.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Brain, C. K. 1967. \u201cThe Transvaal Museum's Fossil Project at Swartkrans.\u201d <em>South African Journal of Science<\/em> 63 (9): 378\u2013384.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1938a. \u201cMore Discoveries of Australopithecus.\u201d <em>Nature<\/em> 141 (1): 828\u2013829.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1938b. \u201cThe Pleistocene Anthropoid Apes of South Africa.\u201d <em>Nature<\/em> 142 (3591): 377\u2013379.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1947. \u201cDiscovery of a New Skull of the South African Ape-Man, Plesianthropus.\u201d <em>Nature<\/em> 159 (4046): 672.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1950. \u201cThe Genera and Species of the South African Fossil Ape-Man.\u201d <em>American Journal of Physical Anthropology<\/em> 8 (1): 1\u201314.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Brunet, Michel, Alain Beauvilain, Yves Coppens, Emile Heintz, Aladji HE Moutaye, and David Pilbeam. 1995. \u201cThe First Australopithecine 2,500 Kilometers West of the Rift Valley (Chad).\u201d <em>Nature<\/em> 378 (6554): 275\u2013273.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Cerling, Thure E., Jonathan G. Wynn, Samuel A. Andanje, Michael I. Bird, David Kimutai Korir, Naomi E. Levin, William Mace, Anthony N. Macharia, Jay Quade, and Christopher H. Remien. 2011. \u201cWoody Cover and Hominin Environments in the Past 6 Million Years.\u201d <em>Nature<\/em> 476, no. 7358 (2011): 51-56..<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clarke, Ronald J. 1998. \u201cFirst Ever Discovery of a Well-Preserved Skull and Associated Skeleton of <em>Australopithecus<\/em>.\u201d <em>South African Journal of Science<\/em> 94 (10): 460\u2013463.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Clarke, Ronald J. 2013. \u201cAustralopithecus from Sterkfontein Caves, South Africa.\u201d In <em>The Paleobiology of Australopithecus<\/em>, edited by K. E. Reed, J. G. Fleagle, and R. E. Leakey, 105\u2013123. Netherlands: Springer.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clarke, Ronald J., and Kathleen Kuman. 2019. \u201cThe Skull of StW 573, a 3.67 Ma Australopithecus Prometheus Skeleton from Sterkfontein Caves, South Africa.\u201d <em>Journal of <\/em><em>H<\/em><em>uman <\/em><em>E<\/em><em>volution<\/em> 134: 102634.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clarke, R. J., and P. V. Tobias. 1995. \u201cSterkfontein Member 2 Foot Bones of the Oldest South African Hominid.\u201d <em>Science<\/em> 269 (5223): 521\u2013524.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Constantino, P. J., and B. A. Wood. 2004. \u201cParanthropus Paleobiology\u201d. In <em>Miscelanea en <\/em><em>Homenae<\/em><em> a Emiliano Aguirre<\/em><em>,<\/em> <em>v<\/em><em>olumen III: Paleoantropologia<\/em>, edited by E. G. P\u00e9rez and S. R. Jara, 136\u2013151. Alcal\u00e1 de Henares: Museo Arqueologico Regional.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Constantino, P. J., and B. A. Wood. 2007. \u201cThe Evolution of Zinjanthropus boisei.\u201d <em>Evolutionary Anthropology: <\/em><em>Issues, News, and Reviews<\/em> 16 (2): 49\u201362.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Dart, Raymond A. 1925. \u201cAustralopithecus africanus, the Man-Ape of South Africa.\u201d <em>Nature<\/em> 115: 195\u2013199.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Darwin, Charles. 1871. <em>The Descent of Man: And Selection in Relation to Sex<\/em>. London: J. Murray.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Daver, Guillaume, F. Guy, Hassane Ta\u00efsso Mackaye, Andossa Likius, J-R. Boisserie, Abderamane Moussa, Laurent Pallas, Patrick Vignaud, and N\u00e9koulnang D. Clarisse. 2022. \"Postcranial Evidence of Late Miocene Hominin Bipedalism in Chad.\" <em>Nature<\/em> 609 (7925): 94\u2013100.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Heinzelin, Jean de, J. Desmond Clark, Tim White, William Hart, Paul Renne, Giday WoldeGabriel, Yonas Beyene, and Elisabeth Vrba. 1999. \u201cEnvironment and Behavior of 2.5-Million-Year-Old Bouri Hominids.\u201d <em>Science<\/em> 284 (5414): 625\u2013629.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">DeMenocal, Peter B. D. 2004. \u201cAfrican Climate Change and Faunal Evolution during the Pliocene\u2013Pleistocene.\u201d <em>Earth and Planetary Science Letters<\/em> 220 (1\u20132): 3\u201324.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">DeMenocal, Peter B. D. and J. Bloemendal, J. 1995. \u201cPlio-Pleistocene Climatic Variability in Subtropical Africa and the Paleoenvironment of Hominid Evolution: A Combined Data-Model Approach.\u201d In <em>Paleoclimate and Evolution, with Emphasis on Human Origins<\/em>, edited by E. S. Vrba, G. H. Denton, T. C. Partridge, and L. H. Burckle, 262\u2013288. New Haven: Yale University Press.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Dirks, Paul HGM, Job M. Kibii, Brian F. Kuhn, Christine Steininger, Steven E. Churchill, Jan D. Kramers, Robyn Pickering, Daniel L. Farber, Anne-Sophie M\u00e9riaux, Andy I. R. Herries, Geoffrey C. P. King, And Lee R. Berger. 2010. \u201cGeological Setting and Age of <em>Australopithecus sediba<\/em> from Southern Africa.\u201d <em>Science<\/em> 328 (5975): 205\u2013208.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Faith, J. Tyler, and Anna K. Behrensmeyer. 2013. \u201cClimate Change and Faunal Turnover: Testing the Mechanics of the Turnover-Pulse Hypothesis with South African Fossil Data.\u201d <em>Paleobiology<\/em> 39 (4): 609\u2013627.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Grine, Frederick E. 1988. \u201cNew Craniodental Fossils of <em>Paranthropus<\/em> from the Swartkrans Formation and Their Significance in \u2018Robust\u2019 Australopithecine Evolution.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by F. E. Grine, 223\u2013243. New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Grine, Frederick E., Carrie S. Mongle, John G. Fleagle, and Ashley S. Hammond. 2022. \"The Taxonomic Attribution of African Hominin Postcrania from the Miocene through the Pleistocene: Associations and Assumptions.\" <em>Journal of Human Evolution<\/em> 173: 103255.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Haile-Selassie, Yohannes, Luis Gibert, Stephanie M. Melillo, Timothy M. Ryan, Mulugeta Alene, Alan Deino, Naomi E. Levin, Gary Scott, and Beverly Z. Saylor. 2015. \u201cNew Species from Ethiopia Further Expands Middle Pliocene Hominin Diversity.\u201d <em>Nature<\/em> 521 (7553): 432\u2013433.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Haile-Selassie, Yohannes, Stephanie M. Melillo, Antonino Vazzana, Stefano Benazzi, and Timothy M. Ryan. 2019. \u201cA 3.8-Million-Year-Old Hominin Cranium from Woranso-Mille, Ethiopia.\u201d <em>Nature<\/em> 573 (7773): 214-219.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Harmand, Sonia, Jason E. Lewis, Craig S. Feibel, Christopher J. Lepre, Sandrine Prat, Arnaud Lenoble, Xavier Bo\u00ebs et al. 2015. \u201c3.3-Million-Year-Old Stone Tools from Lomekwi3, West Turkana, Kenya.\u201d <em>Nature<\/em> 521 (7552): 310\u2013316.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Hay, Richard L. 1990. \u201cOlduvai Gorge: A Case History in the Interpretation of Hominid Paleoenvironments.\u201d In <em>East Africa: Establishment of a Geologic Framework for Paleoanthropology<\/em>, edited by L. Laporte, 23\u201337<em>.<\/em> Boulder: Geological Society of America.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Hay, Richard L., and Mary D. Leakey. 1982. \u201cThe Fossil Footprints of Laetoli.\u201d <em>Scientific American<\/em> 246 (2): 50\u201357.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Hlazo, Nomawethu. 2015. \u201cParanthropus: Variation in Cranial Morphology.\u201d Honours thesis, Archaeology Department, University of Cape Town, Cape Town.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Hlazo, Nomawethu. 2018. \u201cVariation and the Evolutionary Drivers of Diversity in the Genus <em>Paranthropus<\/em>.\u201d Master\u2019s thesis, Archaeology Department, University of Cape Town, Cape Town.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Johanson, D. C., T. D. White, and Y. Coppens. 1978. \u201cA New Species of the Genus <em>Australopithecus<\/em> (Primates: Hominidae) from the Pliocene of East Africa.\u201d <em>Kirtlandia<\/em> 28: 1\u201314.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kimbel, William H. 2015. \u201cThe Species and Diversity of Australopiths.\u201d In <em>Handbook of Paleoanthropology<\/em>, 2nd ed., edited by T. Hardt, 2071\u20132105. Berlin: Springer.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kimbel, William H., and Lucas K. Delezene. 2009. \u201c\u2018Lucy\u2019 Redux: A Review of Research on <em>Australopithecus afarensis<\/em>.\u201d <em>American <\/em><em>J<\/em><em>ournal of <\/em><em>P<\/em><em>hysical <\/em><em>A<\/em><em>nthropology<\/em> 140 (S49): 2\u201348.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Kingston, John D. 2007. \u201cShifting Adaptive Landscapes: Progress and Challenges in Reconstructing Early Hominid Environments.\u201d <em>American Journal of Physical Anthropology<\/em> 134 (S45): 20\u201358.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Kingston, John D., and Terry Harrison. 2007. \u201cIsotopic Dietary Reconstructions of Pliocene Herbivores at Laetoli: Implications for Early Hominin Paleoecology.\u201d <em>Palaeogeography, Palaeoclimatology, Palaeoecology<\/em> 243 (3\u20134): 272\u2013306.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Leakey, Louis S. B. 1959. \u201cA New Fossil Skull from Olduvai.\u201d <em>Nature<\/em> 184 (4685): 491\u2013493.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Leakey, Mary 1971. <em>Olduvai Gorge<\/em>, Vol. 3. Cambridge: Cambridge University Press.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Leakey, Mary D., and Richard L. Hay. 1979. \u201cPliocene Footprints in the Laetoli Beds at Laetoli, Northern Tanzania.\u201d <em>Nature<\/em> 278 (5702): 317\u2013323.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Leakey, Meave G., Craig S. Feibel, Ian McDougall, and Alan Walker. 1995. \u201cNew Four\u2013Million-Year-Old Hominid Species from Kanapoi and Allia Bay, Kenya.\u201d <em>Nature<\/em> 376 (6541): 565\u2013571.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Meave G., Fred Spoor, Frank H. Brown, Patrick N. Gathogo, Christopher Kiarie, Louise N. Leakey, and Ian McDougall. 2001. \u201cNew Hominin Genus from Eastern Africa Shows Diverse Middle Pliocene Lineages.\u201d <em>Nature<\/em> 410 (6827): 433\u2013440.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lebatard, Anne-Elisabeth, Didier L. Bourl\u00e8s, Philippe Duringer, Marc Jolivet, R\u00e9gis Braucher, Julien Carcaillet, Mathieu Schuster et al. 2008. \u201cCosmogenic Nuclide Dating of <em>Sahelanthropus tchadensis<\/em> and <em>Australopithecus bahrelghazali<\/em>: Mio-Pliocene Hominids from Chad.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 105 (9): 3226\u20133231.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lee-Thorp, Julia. 2011. \u201cThe Demise of \u2018Nutcracker Man.\u2019\u201d <em>Proceedings of the National Academy of Sciences<\/em> 108 (23): 9319\u20139320.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lombard, Marlize, L. Y. N. Wadley, Janette Deacon, Sarah Wurz, Isabelle Parsons, Moleboheng Mohapi, Joane Swart, and Peter Mitchell. 2012. \u201cSouth African and Lesotho Stone Age Sequence Updated.\u201d <em>The South African Archaeological Bulletin<\/em> 67 (195): 123\u2013144.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Maslin, Mark A., Chris M. Brierley, Alice M. Milner, Susanne Shultz, Martin H. Trauth, and Katy E. Wilson. 2014. \u201cEast African Climate Pulses and Early Human Evolution.\u201d <em>Quaternary Science Reviews<\/em> 101: 1\u201317.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">McHenry, Henry M. 2009. \u201cHuman Evolution.\u201d In <em>Evolution: The First Four Billion Years<\/em>, edited by M. Ruse and J. Travis, 256\u2013280. Cambridge: The Belknap Press of Harvard University Press..<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Patterson, Bryan, and William W. Howells. 1967. \u201cHominid Humeral Fragment from Early Pleistocene of Northwestern Kenya.\u201d <em>Science<\/em> 156 (3771): 64\u201366.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Pickering, Robyn, and Jan D. Kramers. 2010. \u201cRe-appraisal of the Stratigraphy and Determination of New U-Pb Dates for the Sterkfontein Hominin Site.\u201d <em>Journal of Human Evoluti<\/em><em>on<\/em> 59 (1): 70\u201386.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Potts, Richard. 1998. \u201cEnvironmental Hypotheses of Hominin Evolution.\u201d <em>American Journal of Physical Anthropology<\/em> 107 (S27): 93\u2013136.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Potts, Richard. 2013. \u201cHominin Evolution in Settings of Strong Environmental Variability.\u201d <em>Quaternary Science Reviews<\/em> 73: 1\u201313.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Rak, Yoel. 1983. <em>The <\/em><em>A<\/em><em>ustralopithecine <\/em><em>F<\/em><em>ace<\/em>. New York: Academic Press.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Rak, Yoel. 1988. \u201cOn Variation in the Masticatory System of <em>Australopithecus boisei<\/em>.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by M. Ruse and J. Travis, 193\u2013198<em>.<\/em> New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Semaw, Sileshi. 2000. \u201cThe World\u2019s Oldest Stone Artefacts from Gona, Ethiopia: Their Implications for Understanding Stone Technology and Patterns of Human Evolution between 2.6 Million Years Ago and 1.5 Million Years Ago.\u201d <em>Journal of Archaeological Science<\/em> 27(12): 1197\u20131214.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Shipman, Pat. 2002. <em>The Man Who Found the Missing Link: Eug<\/em><em>e<\/em><em>ne Dubois and <\/em><em>h<\/em><em>is Lifelong Quest to Prove Darwin Right<\/em>. New York: Simon &amp; Schuster.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Spoor, Fred. 2015. \u201cPalaeoanthropology: The Middle Pliocene Gets Crowded.\u201d<em> Nature<\/em> 521 (7553): 432\u2013433.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Strait, David S., Frederick E. Grine, and Marc A. Moniz. 1997. A Reappraisal of Early Hominid Phylogeny.\u201d <em>Journal of <\/em><em>H<\/em><em>uman <\/em><em>E<\/em><em>volution<\/em> 32 (1): 17\u201382.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Thackeray, J. Francis. 2000. \u201c\u2018Mrs. Ples\u2019 from Sterkfontein: Small Male or Large Female?\u201d <em>The South African Archaeological <\/em><em>Bulletin<\/em> 55: 155\u2013158.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Thackeray, J. Francis, Jos\u00e9 Braga, Jacques Treil, N. Niksch, and J. H. Labuschagne. 2002. \u201c\u2018Mrs. Ples\u2019 (Sts 5) from Sterkfontein: An Adolescent Male?\u201d <em>South African Journal of Science<\/em> 98 (1\u20132): 21\u201322.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Toth, Nicholas. 1985. \u201cThe Oldowan Reassessed.\u201d <em>Journal of Archaeological Science<\/em>\u00a012 (2): 101\u2013120.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Vrba, E. S. 1988. \u201cLate Pliocene Climatic Events and Hominid Evolution.\u201d In <em>The <\/em><em>E<\/em><em>volutionary <\/em><em>H<\/em><em>istory of the <\/em><em>R<\/em><em>obust Australopithecines<\/em>, edited by F. E. Grine, 405\u2013426. New York: Aldine.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Vrba, Elisabeth S. 1998. \u201cMultiphasic Growth Models and the Evolution of Prolonged Growth Exemplified by Human Brain Evolution.\u201d <em>Journal of Theoretical Biology<\/em> 190 (3): 227\u2013239.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Vrba, Elisabeth S. 2000. \u201cMajor Features of Neogene Mammalian Evolution in Africa.\u201d In <em>Cenozoic <\/em><em>G<\/em><em>eology of <\/em><em>S<\/em><em>outhern Africa<\/em>, edited by T. C. Partridge and R. Maud, 277\u2013304<em>.<\/em> Oxford: Oxford University Press.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Walker, Alan C., and Richard E. Leakey. 1988. \u201cThe Evolution of <em>Australopithecus boisei<\/em>.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by F. E. Grine, 247\u2013258. New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Walker, Alan, Richard E. Leakey, John M. Harris, and Francis H. Brown. 1986. \u201c2.5-my <em>Australopithecus boisei<\/em> from West of Lake Turkana, Kenya.\u201d <em>Nature<\/em> 322 (6079): 517\u2013522.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Ward, Carol, Meave Leakey, and Alan Walker. 1999. \u201cThe New Hominid Species <em>Australopithecus anamensis<\/em>.\u201d <em>Evolutionary Anthropology<\/em> 7 (6): 197\u2013205.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">White, Tim D. 1988. \u201cThe Comparative Biology of \u2018Robust\u2019 Australopithecus: Clues from Content.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by F. E. Grine, 449\u2013483. New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">White, Tim D., Gen Suwa, and Berhane Asfaw. 1994. \u201c<em>Australopithecus ramidus<\/em>, a New Species of Early Hominid from Aramis, Ethiopia.\u201d <em>Nature<\/em> 371 (6495): 306\u2013312.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Wood, Bernard. 2010. \u201cReconstructing Human Evolution: Achievements, Challenges, and Opportunities.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 10 (2): 8902\u20138909.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Wood, Bernard, and Eve K. Boyle. 2016. \u201cHominin Taxic Diversity: Fact or Fantasy?\u201d <em>Yearbook of Physical Anthropology<\/em> 159 (S61): 37\u201378.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Wood, Bernard, and Kes Schroer. 2017. \u201cParanthropus: Where Do Things Stand?\u201d In <em>Human Paleontology and Prehistory<\/em>, edited by A. Marom and E. Hovers, 95\u2013107. New York: Springer, Cham.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Acknowledgements<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">All of the authors in this section are students and early career researchers in paleoanthropology and related fields in South Africa (or at least have worked in South Africa). We wish to thank everyone who supports young and diverse talent in this field and would love to further acknowledge Black, African, and female academics who have helped pave the way for us.<\/span><\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1777\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1777\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\">Jonathan Marks, Ph.D., University of North Carolina at Charlotte<\/p>\n<p class=\"import-Normal\">Adam P. Johnson, M.A., University of North Carolina at Charlotte\/University of Texas at San Antonio<\/p>\n<p class=\"import-Normal\"><em>This chapter is an adaptation of \"<\/em><a class=\"rId9\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__\/\"><em>Chapter 2: Evolution<\/em><\/a><em>\u201d by Jonathan Marks. In <\/em><a class=\"rId10\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, first edition<\/em><\/a><em>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under <\/em><a class=\"rId11\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\"><em>CC BY-NC 4.0<\/em><\/a><em>. <\/em><\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\"><span style=\"color: #ffffff\">Learning Objectives<\/span><\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li>Explain the relationship among genes, bodies, and organismal change.<\/li>\n<li>Discuss the shortcomings of simplistic understandings of genetics.<\/li>\n<li>Describe what is meant by the \"biopolitics of heredity.\"<\/li>\n<li>Discuss issues caused by misuse of ideas about adaptations and natural selection.<\/li>\n<li>Examine and correct myths about evolution.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p class=\"import-Normal\">The Human Genome Project, an international initiative launched in 1990, sought to identify the entire genetic makeup of our species. For many scientists, it meant trying to understand the genetic underpinnings of what made humans uniquely human. James Watson, a codiscoverer of the helical shape of DNA, wrote that \u201cwhen finally interpreted, the genetic messages encoded within our DNA molecules will provide the ultimate answers to the chemical underpinnings of human existence\u201d (Watson 1990, 248). The underlying message is that what makes humans unique can be found in our <strong>genes<\/strong>. The Human Genome Project hoped to find the core of who we are and where we come from.<\/p>\n<p class=\"import-Normal\">Despite its lofty goal, the Human Genome Project\u2014even after publishing the entire human genome in January 2022\u2014could not fully account for the many factors that contribute to what it is to be human. Richard Lewontin, Steven Rose, and Leon Kamin (2017) argue that genetic determinism of the sort assumed by the Human Genome Project neglects other essential dimensions that contribute to the development and evolution of human bodies, not to mention the role that culture plays. They use an apt metaphor of a cake to illustrate the incompleteness of reductive models. Consider the flavor of a cake and think of the ingredients listed in the recipe. The recipe includes ingredients such as flour, sugar, shortening, vanilla extract, eggs, and milk. Does raw flour taste like cake? Does sugar, vanilla extract, or any of the other ingredients taste like cake? They do not, and knowing the individual flavors of each ingredient does not tell us much about what cake tastes like. Even mixing all of the ingredients in the correct proportions does not get us cake. Instead, external factors such as baking at the right temperature, for the right amount of time, and even the particularities of our evolved sense of taste and smell are all necessary components of experiencing the cake.<\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ff00ff\">Lewontin, Rose, and Kamin (2017) argue that the same is true for humans and other organisms.<\/span><\/p>\n<p class=\"import-Normal\">Knowing everything about cake ingredients does not allow us to fully know cake. Equally so, knowing everything about the genes found in our DNA does not allow us to fully know humans. Different, interacting levels are implicated in the development and evolution of all organisms, including humans. Genes, the structure of chromosomes, developmental processes, epigenetic tags, environmental factors, and still-other components all play key roles such that genetically reductive models of human development and evolution are woefully inadequate.<\/p>\n<p class=\"import-Normal\">The complex interactions across many levels\u2014genetic, developmental, and environmental\u2014explain why we still do not know how our one-dimensional DNA nucleotide sequence results in a four-dimensional organism. This was the unfulfilled promise of the inception of the Human Genome Project in the 1980s and 1990s: the project produced the complete DNA sequence of a human cell in the hopes that it would reveal how human bodies are built and how to cure them when they are built poorly. Yet, that information has remained elusive. Presumably, the knowledge of how organisms are produced from DNA sequences will one day permit us to reconcile the discrepancies between patterns in anatomical evolution and molecular evolution.<\/p>\n<p class=\"import-Normal\">In this chapter, we will consider multilevel evolution and explore evolution as a complex interaction between genetic and epigenetic factors as well as the environments in which organisms live. Next, we will examine the biopolitical nature of human evolution. We will then investigate problems that arise from attributing all traits to an adaptive function. Finally, we will address common misconceptions about evolution. The goal of this chapter is to provide you with the necessary toolkit for understanding the molecular, anatomical, and political dimensions of evolution.<\/p>\n<h2 class=\"import-Normal\">Evolution Happens at Multiple Levels<\/h2>\n<p class=\"import-Normal\">Following Richard Dawkins\u2019s publication of <em>The Selfish Gene <\/em>in 1976, the scientific imagination was captured by the potential of genomics to reveal how genes are copied by Darwinian selection. Dawkins argues that the genes in individuals that contribute to greater reproductive success are the units of selection. His conception of evolution at the molecular level undercuts the complex interactions between organisms and their environments, which are not expressed genomically but are nevertheless key drivers in evolution.<\/p>\n<p class=\"import-Normal\">By the 1980s, the acknowledgment among most biologists that even though genes construct bodies, genes and bodies evolve at different rates and with distinct patterns. This realization led to a renewed focus on how bodies change. The Evolutionary Synthesis of the 1930s\u20131970s had reduced organisms to their <strong>genotypes<\/strong> and species to their <strong>gene pools<\/strong>, which provided valuable insights about the processes of biological change, but it was only a first approximation. Animals are in fact reactive and adaptable beings, not passive and inert genotypes. Species are clusters of socially interacting and reproductively compatible organisms.<\/p>\n<figure style=\"width: 291px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/06\/image8-5.png\" alt=\"An asteroid hits the ocean. Pterodactyls fly among clouds in the foreground.\" width=\"291\" height=\"233\" \/><figcaption class=\"wp-caption-text\">Figure 17.1: A painting by Donald E. Davis representing the Chicxulub asteroid impact off the Yucatan Peninsula that contributed to the mass extinction that included the dinosaurs about 65 million years ago. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Chicxulub_impact_-_artist_impression.jpg\">Chicxulub impact - artist impression<\/a> by Donald E. Davis, <a href=\"https:\/\/www.nasa.gov\/\">NASA<\/a>, is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Once we accept that evolutionary change is fundamentally genetic change, we can ask: How do bodies function and evolve? How do groups of animals come to see one another as potential mates or competitors for mates, as opposed to just other creatures in the environment? Are there evolutionary processes that are not explicable by population genetics? These questions\u2014which lead us beyond reductive assumptions\u2014were raised in the 1980s by Stephen Jay Gould, the leading evolutionary biologist of the late 20th century (see: Gould 2003; 1996).<\/p>\n<p class=\"import-Normal\">Gould spearheaded a movement to identify and examine higher-order processes and features of evolution that were not adequately explained by population genetics. For example, <strong>extinction<\/strong>, which was such a problem for biologists of the 1600s, could now be seen as playing a more complex role in the history of life than population genetics had been able to model. Gould recognized that there are two kinds of extinctions, each with different consequences: background extinctions and mass extinctions. Background extinctions are those that reflect the balance of nature, because in a competitive Darwinian world, some things go extinct and other things take their place. Ecologically, your species may be adapted to its niche, but if another species comes along that\u2019s better adapted to the same niche, eventually your species will go extinct. It sucks, but it is the way of all life: you come into existence, you endure, and you pass out of existence. But mass extinctions are quite different. They reflect not so much the balance of nature as the wholesale disruption of nature: many species from many different lineages dying off at roughly the same time\u2014presumably as the result of some kind of rare ecological disaster. The situation may not be survival of the fittest as much as survival of the luckiest. The result, then, would be an ecological scramble among the survivors. Having made it through the worst, the survivors could now simply divide up the new ecosystem amongst themselves, since their competitors were gone. Something like this may well have happened about 65 million years ago, when a huge asteroid hit the Yucatan Peninsula, which mammals survived but dinosaurs did not (Figure 17.1). Something like this may be happening now, due to human expansion and environmental degradation. Note, though, that there is only a limited descriptive role here for population genetics: the phenomena we are describing are about organisms and species in ecosystems.<\/p>\n<p class=\"import-Normal\">Another question involved the disconnect between properties of <em>species<\/em> and the properties of <em>gene pools<\/em>. For example, there are upwards of 15 species of gibbons but only two species of chimpanzees. Why? There are upwards of 20 species of guenons but fewer than ten of baboons. Why? Are there genes for that? It seems unlikely. Gould suggested that species, as units of nature, might have properties that are not reducible to the genes in their cells. For example, rates of speciation and extinction might be properties of their ecologies and histories rather than their genes. Thus, relationships between environmental contexts and variability within a species result in degrees of resistance to extinction and affect the frequency and rates at which clades diversify (Lloyd and Gould 1993). Consistent biases of speciation rates might well produce patterns of macroevolutionary diversity that are difficult to explain genetically and better understood ecologically. Gould called such biases in speciation rates <strong>species selection<\/strong>\u2014a higher-order process that invokes competition between species, in addition to the classic Darwinian competition between individuals.<\/p>\n<p class=\"import-Normal\">One of Gould\u2019s most important studies involved the very nature of species. In the classical view, a species is continually adapting to its environment until it changes so much that it is a different species than it was at the beginning of this sentence (Eldredge and Gould 1972). That implies that the species is a fundamentally unstable entity through time, continuously changing to fit in. But suppose, argued Gould along with paleontologist Niles Eldredge, a species is more stable through time and only really adapts during periods of ecological instability and change. Then we might expect to find in the fossil record long equilibrium periods\u2014a few million years or so\u2014in which species don\u2019t seem to change much, punctuated by relatively brief periods in which they change a bit and then stabilize again as new species. They called this idea <strong>punctuated equilibria<\/strong>. The idea helps to explain certain features of the fossil record, notably the existence of small anatomical \u201cgaps\u201d between closely related fossil forms (Figure 17.2). Its significance lies in the fact that although it incorporates genetics, punctuated equilibria is not really a theory of genetics but one of types bodies in deep time.<\/p>\n<p class=\"import-Normal\">Punctuated equilibria is seen across taxa, with long periods in the fossil record representing little phenotypic change. These periods of stability are disrupted by shorter periods of rapid <strong>adaptation<\/strong>, the process through which populations of organisms become suited to living in their environments. Phenotypic changes are often coupled with drastic climatic or ecological changes that affect the milieu in which organisms live. For example, throughout much of hominin evolutionary history, brain size was closely associated with body size and thus remained mostly stable. However, changes occurred in average hominin brain size at around 100 thousand years ago, 1 million years ago, and 1.8 million years ago. Several hypotheses have been put forth to explain these changes, including unpredictability in climate and environment (Potts 1998), social development (Barton 1996), and the evolution of language (Deacon 1998). Evidence from the fossil record, paleoclimate models, and comparative anatomy suggests that the changes observed in hominin lineage result from biocultural processes\u2014that is, the coalescence of environmental and cultural factors that selected for larger brains (Marks 2015; Shultz, Nelson, and Dunbar 2012).<\/p>\n<figure style=\"width: 461px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image5-8.png\" alt=\"Two graphs contrast phyletic gradualism and punctuated equilibria.\" width=\"461\" height=\"222\" \/><figcaption class=\"wp-caption-text\">Figure 17.2: Different ways of conceptualizing the evolutionary relationship between an earlier and a later species. With phyletic gradualism, species are envisioned transforming continually in a direct line over time. With punctuated equilibria species branch off at particular points over time.\u00a0 Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__\/\">Phyletic gradualism vs. punctuated equilibria (Figure 2.12)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">In response to the call for a theory of the evolution of form, the field of <strong>evo-devo<\/strong>\u2014the intersection of evolutionary and developmental biology\u2014arose. The central focus here is on how changes in form and shape arise. An embryo matures by the stimulation of certain cells to divide, forming growth fields. The interactions and relationships among these growth fields generate the structures of the body. The <strong>hox genes<\/strong> that regulate these growth fields turn out to be highly conserved across the animal kingdom. This is because they repeatedly turn on and off the most basic genes guiding the animal\u2019s development, and thus any changes to them would be catastrophic. Indeed, these genes were first identified by manipulating them in fruit flies, such that one could produce a bizarre mutant fruit fly that grew a pair of legs where its antennae were supposed to be (Kaufman, Seeger, and Olsen 1990).<\/p>\n<p class=\"import-Normal\">Certain genetic changes can alter the fates of cells and the body parts, while other genetic changes can simply affect the rates at which neighboring groups of cells grow and divide, thus producing physical bumps or dents in the developing body. The result of altering the relationships among these fields of cellular proliferation in the growing embryo is <strong>allometry<\/strong>, or the differential growth of body parts. As an animal gets larger\u2014either over the course of its life or over the course of macroevolution\u2014it often has to change shape in order to live at a different size. Many important physiological functions depend on properties of geometric area: the strength of a bone, for example, is proportional to its cross-sectional area. But area is a two-dimensional quality, while growing takes place in three dimensions\u2014as an increase in mass or volume. As an animal expands, its bones necessarily weaken, because volume expands faster than area does. Consequently a bigger animal has more stress on its bones than a smaller animal does and must evolve bones even thicker than they would be by simply scaling the animal up proportionally. In other words, if you expand a mouse to the size of an elephant, it will nevertheless still have much thinner bones than the elephant does. But those giant mouse bones will unfortunately not be adequate to the task. Thus, a giant mouse would have to change aspects of its form to maintain function at a larger size (see Figure 17.3).<\/p>\n<p class=\"import-Normal\"><img class=\"aligncenter\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image6-6.png\" alt=\"Side-view of a mouse skeleton.\" width=\"515\" height=\"252\" \/><\/p>\n<figure style=\"width: 453px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image7-9.png\" alt=\"Side-view of an elephant skeleton.\" width=\"453\" height=\"326\" \/><figcaption class=\"wp-caption-text\">Figure 17.3: Mouse (top) and elephant (bottom) skeletons. Notice the elephant\u2019s bones are more robust when the two animals are the same size. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__\/\">Mouse and elephant skeletons (Figure 2.13)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Physiologically, we would like to know how the body \u201cknows\u201d when to turn on and off the genes that regulate growth to produce a normal animal. Evolutionarily, we would like to know how the body \u201clearns\u201d to alter the genetic on\/off switch (or the genetic \u201cslow down\/speed up\u201d switch) to produce an animal that looks different. Moreover, since organisms differ from one another, we would like to know how the developing body distinguishes a range of normal variation from abnormal variation. And, finally, how does abnormal variation eventually become normal in a descendant species?<\/p>\n<p class=\"import-Normal\">Taking up these questions, Gould invoked the work of a British geneticist named Conrad H. Waddington, who thought about genetics in less reductive ways than his colleagues. Rather than isolate specific DNA sites to analyze their function, Waddington instead studied the inheritance of an organism\u2019s reactivity\u2014its ability to adapt to the circumstances of its life. In a famous experiment, he grew fruit fly eggs in an atmosphere containing ether. Most died, but a few survived somehow by developing a weird physical feature: a second thorax with a second pair of wings. Waddington bred these flies and soon developed a stable line of flies who would reliably develop a second thorax when grown in ether. Then he began to lower the concentration of ether, while continuing to selectively breed the flies that developed the strange appearance. Eventually he had a line of flies that would stably develop the \u201cbithorax\u201d <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1238\"><strong>phenotype<\/strong><\/a>\u2013the suite of traits of an organism\u2013even when there was no ether; it had become the \u201cnew normal.\u201d The flies had genetically assimilated the bithorax condition.<\/p>\n<p class=\"import-Normal\">Waddington was thus able to mimic the <strong>inheritance of acquired characteristics<\/strong>: what had been a trait stimulated by ether a few generations ago was now a normal part of the development of the descendants. Waddington recognized that while he had performed a selection experiment on genetic variants, he had not selected for particular traits. Rather, he helped produce the physiological tendency to develop particular traits when appropriately stimulated. He called that tendency <strong>plasticity<\/strong> and its converse, the tendency to stay the same even under weird environmental circumstances, <strong>canalization.<\/strong> Waddington had initially selected for plasticity, the tendency to develop the bithorax phenotype under weird conditions, and then, later, for canalization, the developmental normalization of that weird physical trait. Although Waddington had high stature in the community of geneticists, evolutionary biologists of the 1950s and 1960s regarded him with suspicion because he was not working within the standard mindset of reductionism, which saw evolution as the spread of genetic variants that coded for favorable traits. Both Waddington and Gould resisted contemporary intellectual paradigms that favored reductive accounts of evolutionary processes. They conceived of evolution as an emergent process in which many external factors (e.g. climate, environment, predation) and internal factors (e.g., genotypes, plasticity, canalization) coalesce to produce the evolutionary trends that we observe in the fossil record and our genome.<\/p>\n<p class=\"import-Normal\">While Gould and Waddington both looked beyond the genome to understand evolution, the Human Genome Project\u2014an international project with the goal of identifying each base pair in the human genome in the 1990s\u2014generated a great deal of public interest in analyzing the human DNA sequence from the standpoint of medical genetics. Some of the rhetoric aimed to sell the public on investing a lot of money and resources in sequencing the human genome in order to show the genetic basis of heritable traits, cure genetic diseases, and learn what it means ultimately to be biologically human. However, the Human Genome Project was not actually able to answer those questions through the use of genetics alone, and thus a broader, more holistic account was required.<\/p>\n<p class=\"import-Normal\">This holistic account came from decades of research in human biology and anthropology, which understood the human body as highly adaptable, dynamic, and emergent. For example, in the early 20th century, anthropologist Franz Boas measured the skulls of immigrants to the U.S., revealing that environmental, not merely genetic, factors affected skull shape. The growing human body adjusts itself to the conditions of life, such as diet, sunshine, high altitude, hard labor, population density, how babies are carried\u2014any and all of which can have subtle but consistent effects upon its development. There can thus be no normal human form, only a context-specific range of human forms.<\/p>\n<p class=\"import-Normal\">However, what the human biologists called human adaptability, evolutionary biologists called developmental plasticity, and evidence quickly began to mount for its cause being <strong>epigenetic <\/strong>modifications to DNA. Epigenetic modifications are changes to how genes are used by the body (as opposed to changes in the DNA sequences; see Chapter 3). Scientific interest shifted from the focus of the Human Genome Project to the ways that bodies are made by evolutionary-developmental processes, including epigenetics. What is meant by \u201cepigenetic modification\u201d? Evolution is about how descendants diverge from their ancestors. Inheritance from parent to offspring is still critical to this process, which occurs through genetic recombination: the pairing of homologous chromosomes and sharing of genetic material during meiosis (see Chapter 3). However, in the 21st century, the link between evolution and inheritance has broadened with a clearer understanding of how environmental and developmental factors shape bodies and the expression of genes, including epigenetic inheritance patterns. While offspring inherit their genes through random assortment during meiosis, environmental factors also shape how genes are used. When these epigenetic modifications occur in germ cells, they can be passed onto offspring. In these cases, there is no change in the DNA sequence but rather in how genes are used by the body due to DNA methylation and the structure of chromosomes due to histone acetylation (see Chapter 3).<\/p>\n<p class=\"import-Normal\">In addition, we now recognize that evolution is affected by two other forms of intergenerational transmission and inheritance (in addition to genetics and epigenetics). These forms include behavioral variation and culture. That is, behavioral information can be transmitted horizontally (intragenerationally), permitting more rapid ways for organisms to adjust to the environment. And, then there is the fourth mode of transmission: the cultural or symbolic mode. It is argued that humans are the only species that horizontally transmits an arbitrary set of rules to govern communication, social interaction, and thought. This shared information is symbolic and has resulted in what we recognize as \u201cculture\u201d: locally emergent worlds of names, words, pictures, classifications, revered pasts, possible futures, spirits, dead ancestors, unborn descendants, in-laws, politeness, taboo, justice, beauty, and story, all accompanied by practices and a material world of tools.<\/p>\n<p class=\"import-Normal\">Consequently our contemporary ideas about evolution see the evolutionary processes as hierarchically organized and not restricted to the differential transmission of DNA sequences into the next generation. While that is indeed a significant part of evolution, the organism and species are nevertheless crucial to understanding how those DNA sequences get transmitted. Further, the transmission of epigenetic, behavioral, and symbolic information play a complex role in perpetuating our genes, bodies, and species. In the case of human evolution, one can readily see that symbolic information and cultural adaptation are far more central to our lives and our survival today than DNA and genetic adaptation. It is thus misleading to think of humans passively occupying an environmental niche. Rather, humans are actively engaged in constructing our own niches, as well as adapting to them and using them to adapt. The complex interplay between a species and its active engagement in creating its own ecology is known as <strong>niche construction<\/strong>. If we understand <strong>natural selection<\/strong>\u2013the process by which populations adapt to their specific environments\u2013as the effects that environmental context has on the reproductive success of organisms, then niche construction is the process through which organisms shape their own selective pressures.<\/p>\n<h2 class=\"import-Normal\">The Biopolitics of Heredity<\/h2>\n<p class=\"import-Normal\">\u201cScience isn\u2019t political\u201d is a sentiment that you have likely heard before. Science is supposed to be about facts and objectivity. It exists, or at least ought to, outside of petty human concerns. However, the sorts of questions we ask as scientists, the problems we choose to study, the categories and concepts we use, who gets to do science, and whose work gets cited are all shaped by culture. Doing science is a political act. This fact is markedly true for human evolution. While it is easier to create intellectual distance between us and fruit flies and viruses, there is no distance when we are studying ourselves. The hardest lesson to learn about human evolution is that it is intensely political. Indeed, to see it from the opposite side, as it were, the history of creationism\u2014the belief that the universe was divinely created around 6,000 years ago\u2014is essentially a history of legal decisions. For instance, in <em>Tennessee v. John T. Scopes<\/em> (1925), a schoolteacher was prosecuted for violating a law in Tennessee that prohibited the teaching of human evolution in public schools, where teachers were required by law to teach creationism.<\/p>\n<p class=\"import-Normal\">More recently, legal decisions aimed at legislating science education have shaped how students are exposed to evolutionary theory. For instance, <em>McLean v. Arkansas<\/em> (1982) dispatched \u201cscientific creationism\u201d by arguing that the imposition of balanced teaching of evolution and creationism in science classes violates the Establishment Clause, separating church and state. Additionally, <em>Kitzmiller v. Dover (Pennsylvania) Area School District<\/em> (2005) dispatched the teaching of \u201cintelligent design\u201d in public school classrooms as it was deemed to not be science. In some cases, people see unbiblical things in evolution, although most Christian theologians are easily able to reconcile science to the Bible. In other cases, people see immoral things in evolution, although there is morality and immorality everywhere. And some people see evolution as an aspect of alt-religion, usurping the authority of science in schools to teach the rejection of the Christian faith, which would be unconstitutional due to the protected separation of church and state.<\/p>\n<p class=\"import-Normal\">Clearly, the position that politics has nothing to do with science is untenable. But is the politics in evolution an aberration or is it somehow embedded in science? In the early 20th century, scientists commonly promoted the view that science and politics were separate: science was seen as a pure activity, only rarely corrupted by politics. And yet as early as World War I, the politics of nationalism made a hero of the German chemist Fritz Haber for inventing poison gas. And during World War II, both German doctors and American physicists, recruited to the war effort, helped to end many civilian lives. Therefore, we can think of the apolitical scientist as a self-serving myth that functions to absolve scientists of responsibility for their politics. The history of science shows how every generation of scientists has used evolutionary theory to rationalize political and moral positions. In the very first generation of evolutionary science, Darwin\u2019s <em>Origin of Species<\/em> (1859) is today far more readable than his <em>Descent of Man<\/em> (1871). The reason is that Darwin consciously purged <em>The Origin of Species<\/em> of any discussion of people. And when he finally got around to talking about people, in <em>The Descent of Man<\/em>, he simply imbued them with the quaint Victorian prejudices of his age, and the result makes you cringe every few pages. There is plenty of politics in there\u2014sexism, racism, and colonialism\u2014because <em>you cannot talk about people apolitically<\/em>.<\/p>\n<p class=\"import-Normal\">One immediate faddish deduction from Darwinism, popularized by Herbert Spencer (1864) as \u201csurvival of the fittest,\u201d held that unfettered competition led to advancement in nature and to human history. Since the poor were purported losers in that struggle, anything that made their lives easier would go against the natural order. This position later came to be known ironically as \u201cSocial Darwinism.\u201d Spencer was challenged by fellow Darwinian Thomas Huxley (1863), who agreed that struggle was the law of the jungle but observed that we don\u2019t live in jungles anymore. The obligation to make lives better for others is a moral, not a natural, fact. We simultaneously inhabit a natural universe of descent from apes and a moral universe of injustice and inequality, and science is not well served by ignoring the latter.<\/p>\n<p class=\"import-Normal\">Concurrently, the German biologist Ernst Haeckel\u2019s 1868 popularization of Darwinism was translated into English a few years later as <em>The History of Creation<\/em>. As we saw earlier, Haeckel was determined to convince his readers that they were descended from apes, even in the absence of fossil evidence attesting to it. When he made non-Europeans into the missing links that connected his readers to the apes, and depicted them as ugly caricatures, he knew precisely what he was doing. Indeed, even when the degrading racial drawings were deleted from the English translation of his book, the text nevertheless made his arguments quite clear. And a generation later, when the Americans had not yet entered the Great War in 1916, a biologist named Vernon Kellogg visited the German High Command as a neutral observer and found that the officers knew a lot about evolutionary biology, which they had gotten from Haeckel and which rationalized their military aggressions. Kellogg went home and wrote a bestseller about it, called <em>Headquarters Nights<\/em> (1917). World War I would have been fought with or without evolutionary theory, but as a source of scientific authority, evolution\u2014even if a perversion of the Darwinian theory\u2014had very quickly attained global geopolitical relevance.<\/p>\n<p class=\"import-Normal\">Oftentimes, politics in evolutionary science is subtle, due to the pervasive belief in the advancement of science. We recognize the biases of our academic ancestors and modify our scientific stories accordingly. But we can never be free of our own cultural biases, which are invisible to us, as much as our predecessors\u2019 biases were invisible to them. In some cases, the most important cultural issues resurface in different guises each generation, like scientific racism. <strong>Scientific racism<\/strong> is the recruitment of science for the evil political ends of racism, and it has proved remarkably impervious to evolution. Before Darwin, there was creationist scientific racism, and after Darwin, there was evolutionist scientific racism. And there is still scientific racism today, self-justified by recourse to evolution, which means that scientists have to be politically astute and sensitive to the uses of their work to counter these social tendencies.<\/p>\n<p class=\"import-Normal\">Consider this: Are you just your ancestry, or can you transcend it? If that sounds like a weird question, it was actually quite important to a turn-of-the-20th-century European society in which an old hereditary aristocracy was under increasing threat from a rising middle class. And that is why the very first English textbook of Mendelian genetics concluded with the thought that \u201cpermanent progress is a question of breeding rather than of pedagogics; a matter of gametes, not of training \u2026 the creature is not made but born\u201d (Punnett 1905, 60). <em>Translation: Not only do we now know a bit about how heredity works, but it\u2019s also the most important thing about you. Trust me, I\u2019m a scientist.<\/em><\/p>\n<p class=\"import-Normal\">Yet evolution is about how descendants come to differ from ancestors. Do we really know that your heredity, your DNA, your ancestry, is the most important thing about you? That you were born, not made? After all, we do know that you could be born into slavery or as a peasant, and come from a long line of enslaved people or peasants, and yet not have slavery or peasantry be the most important thing about you. Whatever your ancestors were may unfortunately constrain what you can become, but as a moral precept, it should not. But just as science is not purely \u201cfacts and objectivity,\u201d ancestry is not a strictly biological concept. Human ancestry is biopolitics, not biology.<\/p>\n<p class=\"import-Normal\">Evolution is fundamentally a theory about ancestry, and yet ancestors are, in the broad anthropological sense, sacred: ancestors are often more meaningful symbolically than biologically. Just a few years after <em>The Origin of Species <\/em>(Darwin 1859), the British politician and writer Benjamin Disraeli declared himself to be on the side of the angels, not the apes, and to \u201crepudiate with indignation and abhorrence those new-fangled theories\u201d (Monypenny, Flavelle, and Buckle 1920, 105). He turned his back on an ape ancestry and looked to the angel; yet, he did so as a prominent Jew-turned-Anglican, who had personally transcended his humble roots and risen to the pinnacle of the Empire. Ancestry was certainly important, and Disraeli was famously proud of his, but it was also certainly not the most important thing, not the primary determinant of his place in the world. Indeed, quite the opposite: Disraeli\u2019s life was built on the transcendence of many centuries of Jewish poverty and oppression in Europe. Humble ancestry was there to be superseded and nobility was there to be earned; Disraeli would later become the Earl of Beaconsfield. Clearly, \u201care you just your ancestry\u201d is not a value-neutral question, and \u201cthe creature is not made, but born\u201d is not a value-neutral answer.<\/p>\n<p class=\"import-Normal\">Ancestry being the most important thing about a person became a popular idea twice in 20th century science. First, at the beginning of the century, when the <strong>eugenics<\/strong> movement in America called attention to \u201cfeeble-minded stocks,\u201d which usually referred to the poor or to immigrants (see Figure 17.4; and see Chapter 2). This movement culminated in Congress restricting the immigration of \u201cfeeble-minded races\u201d (said to include Jews and Italians) in 1924, and the Supreme Court declaring it acceptable for states to sterilize their \u201cfeeble-minded\u201d citizens involuntarily in 1927. After the Nazis picked up and embellished these ideas during World War II, Americans moved swiftly away from them in some contexts (e.g., for most people of European descent) while still strictly adhering in other contexts (e.g., Japanese internment camps and immigration restrictions).<\/p>\n<figure style=\"width: 374px\" class=\"wp-caption alignright\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image4-6.png\" alt=\"Historic photo. People sit in front of a structure with a \u201cEugenic and Health Exhibit&quot; banner.\" width=\"374\" height=\"262\" \/><figcaption class=\"wp-caption-text\">Figure 17.4: Eugenic and Health Exhibit, Fitter Families exhibit, and examination building, Kansas State Free Fair. Credit: <a href=\"https:\/\/www.dnalc.org\/view\/16328-Gallery-14-Eugenics-Exhibit-at-the-Kansas-State-Free-Fair-1920.html\">Gallery 14: Eugenics Exhibit at the Kansas State Free Fair, 1920 ID (ID 16328)<\/a> by <a href=\"https:\/\/www.dnalc.org\/\">Cold Spring Harbor<\/a> (Courtesy American Philosophical Society) is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/3.0\/us\/\">CC BY-NC-ND 3.0 License.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\">Ancestry again became paramount in the drumming up of public support for the Human Genome Project in the 1990s. Public support for sequencing the human genome was encouraged by a popular science campaign that featured books titled <em>The Book of Man <\/em>(Bodmer and McKie 1997), <em>The Human Blueprint <\/em>(Shapiro 1991), and <em>The Code of Codes<\/em> (Kevles and Hood 1993). These books generally promised cures for genetic diseases and a deeper understanding of the human condition. We can certainly identify progress in molecular genetics over the last couple of decades since the human genome was sequenced, but that progress has notably not been accompanied by cures for genetic diseases, nor by deeper understandings of the human condition.<\/p>\n<p class=\"import-Normal\">Even at the most detailed and refined levels of genetic analysis, we still don\u2019t have much of an understanding of the actual basis by which things seem to \u201crun in families.\u201d While the genetic basis of simple, if tragic, genetic diseases have become well-known\u2014such as sickle-cell anemia, cystic fibrosis, and Tay-Sachs\u2019 Disease\u2014we still haven\u2019t found the ostensible genetic basis for traits that are thought to have a strong genetic component. For example, a recent genetic summary found over 12,000 genetic sites that contributed to height yet still explained only about 40-50 percent of the variation in height among European ancestry but no more than 10-20 percent of variation of other ancestries, which we know strongly runs in families (Yengo et al. 2022).<\/p>\n<p class=\"import-Normal\">Partly in reaction to the reductionistic hype of the Human Genome Project, the study of epigenetics has become the subject of great interest. One famous natural experiment involves a Nazi-imposed famine in Holland over the winter of 1944\u20131945. Children born during and shortly after the famine experienced a higher incidence of certain health problems as adults, many decades later. Apparently, certain genes had been down-regulated early in development and remained that way throughout the course of life. Indeed, this modified regulation of the genes in response to the severe environmental conditions may have been passed on to their children.<\/p>\n<p class=\"import-Normal\">Obviously one\u2019s particular genetic constitution may play an important role in one\u2019s life trajectory. But overvaluing that role may have important social and political consequences. In the first place, genotypes are rendered meaningful in a cultural universe. Thus, if you live in a strongly patriarchal society and are born without a Y chromosome (since human males are chromosomally XY and females XX), your genotype will indeed have a strong effect upon your life course. So even though the variation is natural, the consequences are political. The mediating factors are the cultural ideas about how people of different sexes ought to be treated, and the role of the state in permitting certain people to develop and thrive. More broadly, there are implications for public education if variation in intelligence is genetic. There are implications for the legal system if criminality is genetic. There are implications for the justice system if sexual preference, or sexual identity, is genetic. There are implications for the development of sports talent if that is genetic. And yet, even for the human traits that are more straightforward to measure and known to be strongly heritable, the DNA base sequence variation seems to explain little.<\/p>\n<p class=\"import-Normal\">Genetic determinism or <strong>hereditarianism<\/strong> is the idea that \u201cthe creature is made, not born\u201d\u2014or, in a more recent formulation by James Watson, that \u201cour fate is in our genes.\u201d One of the major implications drawn from genetic determinism is that the feature in question must inevitably express itself; therefore, we can\u2019t do anything about it. Therefore, we might as well not fund the social programs designed to ameliorate economic inequality and improve people\u2019s lives, because their courses are fated genetically. And therefore, they don\u2019t deserve better lives.<\/p>\n<p class=\"import-Normal\">All of the \u201ctherefores\u201d in the preceding paragraph are open to debate. What is important is that the argument relies on a very narrow understanding of the role of genetics in human life, and it misdirects the causes of inequality from cultural to natural processes. By contrast, instead of focusing on genes and imagining them to place an invisible limit upon social progress, we can study the ways in which your DNA sequence does <em>not<\/em> limit your capability for self-improvement or fix your place in a social hierarchy. In general, two such avenues exist. First, we can examine the ways in which the human body responds and reacts to environmental variation: human adaptability and plasticity. This line of research began with the anthropometric studies of immigrants by Franz Boas in the early 20th century and has now expanded to incorporate the epigenetic inheritance of modified human DNA. And second, we can consider how human lives are shaped by social histories\u2014especially the structural inequalities within the societies in which they grow up.<\/p>\n<p class=\"import-Normal\">Although it arises and is refuted every generation, the radical hereditarian position (genetic determinism) perennially claims to speak for both science and evolution. It does not. It is the voice of a radical fringe\u2014perhaps naive, perhaps evil. It is not the authentic voice of science or of evolution. Indeed, keeping Charles Darwin\u2019s name unsullied by protecting it from association with bad science often seems like a full-time job. Culture and epigenetics are very much a part of the human condition, and their roles are significant parts of the complete story of human evolution.<\/p>\n<p><span style=\"background-color: #00ffff\"><span style=\"text-decoration: underline\">(Sterilization of Indigenous women in Canada)<\/span> (https:\/\/www.thecanadianencyclopedia.ca\/en\/article\/sterilization-of-indigenous-women-in-canada)\u00a0<\/span><\/p>\n<h2 class=\"import-Normal\">Adaptationism and the Panglossian Paradigm<\/h2>\n<p class=\"import-Normal\">The story of human evolution, and the evolution of all life for that matter, is never settled because evolution is ongoing. Additionally, because the conditions that shape evolutionary trajectories are not predetermined, evolution itself is emergent. Even during periods of ecological stability, when fewer macroevolutionary changes occur, populations of organisms continue to experience change. When ecological stability is disrupted, populations must adapt to the changes. Darwin explained in naturalistic terms how animals adapt to their environments: traits that contribute to an organism's ability to survive and reproduce in specific environments will become more common. The most \u201cfit\u201d\u2014those organisms best suited to the <em>current<\/em> environmental conditions in which they live\u2014have survived over eons of the history of life on earth to cocreate ecosystems full of animals and plants. Our own bodies are full of evident adaptations: eyes for seeing, ears for hearing, feet for walking on, and so forth.<\/p>\n<p class=\"import-Normal\">But what about hands? Feet are adapted to be primarily weight-bearing structures (rather than grasping structures, as in the apes) and that is what we primarily use them for. But we use our hands in many ways: for fine-scale manipulation, greeting, pointing, stimulating a sexual partner, writing, throwing, and cooking, among other uses. So which of these uses express what hands are \u201cfor,\u201d when all of them express what hands do?<\/p>\n<p class=\"import-Normal\">Gould and Lewontin (1979) illustrate the problem with assuming that the function of a trait defines its evolutionary cause. Consider the case of Dr. Pangloss\u2014the protagonistic of Voltaire\u2019s <em>Candide<\/em>\u2014who believed that we lived in the best of all possible worlds. Gould and Lewontin use his pronouncement that \u201cnoses were made for spectacles and so we have spectacles\u201d to demonstrate the problem with assuming any trait has evolved for a specific purpose. Identifying a function of a trait does not necessitate that the function is the ultimate cause of the trait. Individual traits are not under selection pressures in isolation; in fact, an entire organism must be able to survive and reproduce in their environment. When natural selection results in adaptations, changes that occur in some traits can have cascading effects throughout the phenotype and features that are not under selection pressure can also change.<\/p>\n<figure style=\"width: 279px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image3-5.png\" alt=\"Human hand is smaller with smaller fingers and smoother skin compared to a chimpanzee hand.\" width=\"279\" height=\"264\" \/><figcaption class=\"wp-caption-text\">Figure 17.5: Drawings of a human hand (left) and a chimpanzee hand (right). Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__\/\">Human and chimpanzee hand (Figure 2.16)<\/a> by Mary Nelson original to <a href=\"https:\/\/explorations.americananthro.org\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">There is an important lesson in recognizing that what things do in the present is not a good guide to understanding why they came to exist. Gunpowder was invented for entertainment\u2014only later was it adopted for killing people. The Internet was invented to decentralize computers in case of a nuclear attack\u2014and only later adopted for social media. Apes have short thumbs and use their hands in locomotion; our ancestors stopped using their hands in locomotion by about six million years ago and had fairly modern-looking hands by about two million years ago. We can speculate that a combination of selection for abstract thought and dexterity led to evolution of the human hand, with its capability for toolmaking that exceeds what apes can do (see Figure 17.5). But let\u2019s face it\u2014how many tools have you made today?<\/p>\n<p class=\"import-Normal\">Consequently, we are obliged to see the human foot as having a purpose to which it is adapted and the human hand as having multiple purposes, most of which are different from what it originally evolved for. Paleontologists Gould and Elisabeth Vrba suggested that an original use be regarded as an adaptation and any additional uses be called \u201c<strong>exaptations.<\/strong>\u201d Thus, we would consider the human hand to be an adaptation for toolmaking and an exaptation for writing. So how do we know whether any particular feature is an adaptation, like the walking foot, rather than an exaptation, like the writing hand? Or more broadly, how can we reason rigorously from what a feature does to what it evolved for?<\/p>\n<p class=\"import-Normal\">The answer to the question \u201cwhat did this feature evolve for?\u201d creates an origin myth. This origin myth contains three assumptions: (1) features can be isolated as evolutionary units; (2) there is a specific reason for the existence of any particular feature; and (3) there is a clear and simplistic explanation for why the feature evolved.<\/p>\n<figure style=\"width: 378px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image9-8.png\" alt=\"Head with images and human qualities drawn on it. Journal title printed at the bottom.\" width=\"378\" height=\"437\" \/><figcaption class=\"wp-caption-text\">Figure 17.6: According to the early 19th century science of phrenology, units of personality could be mapped onto units in the head, as shown on this cover of The Phrenology Journal. Credit: <a href=\"https:\/\/wellcomecollection.org\/works\/b6skynug\">Phrenology; Chart<\/a> [slide number 5278, photo number: L0000992, original print from Dr. E. Clark, The Phrenological Journal (Know Thyself)] by <a href=\"https:\/\/wellcomecollection.org\/\">Wellcome Collection<\/a>, is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/legalcode\">CC BY 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">The first assumption was appreciated a century ago as the \u201cunit-character problem.\u201d Are the units by which the body grows and evolves the same as units we name? This is clearly not the case: we have genes and we have noses, and we have genes that affect noses, but we don\u2019t have \u201cnose genes.\u201d What is the relationship between the evolving elements that we see, identify, and name, and the elements that biologically exist and evolve? It is hard to know, but we can use the history of science as a guide to see how that fallacy has been used by earlier generations. Back in the 19th century, the early anatomists argued that since the brain contained the mind, they could map different mental states (acquisitiveness, punctuality, sensitivity) onto parts of the brain. Someone who was very introspective, say, would have an enlarged introspection part of the brain, a cranial bulge to represent the hyperactivity of this mental state. The anatomical science was known as <strong>phrenology<\/strong>, and it was predicated on the false assumption that units of thought or personality or behavior could be mapped to distinct parts of the brain and physically observed (see Figure17.6). This is the fallacy of reification, imagining that something named is something real.<\/p>\n<figure style=\"width: 295px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image1-8-1.png\" alt=\"A black-and-white drawing of a chimpanzee head and face.\" width=\"295\" height=\"236\" \/><figcaption class=\"wp-caption-text\">Figure 17.7: Chimpanzees have big ears. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Chimpanzee_head_sketch.png\">Chimpanzee head sketch<\/a> by <a href=\"https:\/\/de.wikipedia.org\/wiki\/Benutzer:Roger_Zenner\">Roger Zenner<\/a>, original by Brehms Tierleben (1887), is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">The second assumption, that everything has a reason, has long been recognized as a core belief of religion. Our desire to impose order and simplicity on the workings of the universe, however, does not constrain it to obey simple and orderly causes. Magic, witchcraft, spirits, and divine agency are all powerful explanations for why things happen. Consequently, it is probably not a good idea to lump natural selection in with those. Sometimes things do happen for a reason, of course, but other times things happen as byproducts of other things, or for very complicated and entangled reasons, or for no reason at all. What phenomena have reasons and thereby merit explanation? Chimpanzees have very large testicles, and we think we know why: their promiscuous sexual behavior triggers intense competition for high sperm count. But chimpanzees also have very large ears, but much less scientific attention has been paid to this trait (see Figure 17.7). Why not? Why should there be a reason for chimp testicles but not for chimp ears? What determines the kinds of features that we try to explain, as opposed to the ones that we do not? Again, the assumption that any specific feature has a reason is metaphysical; that is to say, it may be true in any particular case, but to assume it in all cases is gratuitous.<\/p>\n<p class=\"import-Normal\">And third, the possibility of knowing what the reason for any particular feature is, assuming that it has one, is a challenge for evolutionary epistemology (the theory of how we know things). Consider the big adaptations of our lineage: bipedalism and language. Nobody doubts that they are good, and they evolved by natural selection, and we know how they work. But why did they evolve? If talking and walking are simply better than not talking and not walking, then why did they evolve in just a single branch of the ape lineage in the primate family tree? We don\u2019t know what bipedalism evolved for, although there are plenty of speculations: walking long distances, running long distances, cooling the head, seeing over tall grass, carrying babies, carrying food, wading, threatening, counting calories, sexual display, and so on. Neither do we know what language evolved for, although there are speculations: coordinating hunting, gossiping, manipulating others. But it is also possible that bipedality is simply the way that a small arboreal ape travels on the ground, if it isn\u2019t in the treetops. Or that language is simply the way that a primate with small canine teeth and certain mental propensities comes to communicate. If that were true, then there might be no reason for bipedality or language: having the unique suite of preconditions and a fortuitous set of circumstances simply set them in motion, and natural selection elaborated and explored their potentials. It is possible that walking and talking simply solved problems that no other lineage had ever solved; but even if so, the fact remains that the rest of the species in the history of life have done pretty well without having solved them.<\/p>\n<p class=\"import-Normal\">It is certainly very optimistic to think that all three assumptions (that organisms can be meaningfully atomized, that everything has a reason, and that we can know the reason) would be simultaneously in effect. Indeed, just as there are many ways of adapting (genetically, epigenetically, behaviorally, culturally), there are also many ways of being nonadaptive, which would imply that there is no reason at all for the feature in question.<\/p>\n<p class=\"import-Normal\">First, there is the element of randomness of population histories. There are more cases of sickle-cell anemia among sub-Saharan Africans than other peoples, and there is a reason for it: carriers of sickle-cell anemia have a resistance to malaria, which is more frequent in parts of Africa (as discussed in Chapters 4 and 14). But there are more cases of a blood disease called variegated porphyria, a rare genetic metabolic disorder, in the Afrikaners of South Africa (descendants of mostly Dutch settlers in the 17th century) than in other peoples, and there is no reason for it. Yet we know the cause: One of the founding Dutch colonial settlers had the <strong>allele<\/strong>\u2013a variant of a gene\u2013and everyone in South Africa with it today is her descendant. But that is not a reason\u2014that is simply an accident of history.<\/p>\n<p class=\"import-Normal\">Second, there is the potential mismatch between the past and the present. The value of a particular feature in the past may be changed as the environmental circumstances change. Our species is diurnal, and our ancestors were diurnal. But beginning around a few hundred thousand years ago, our ancestors could build fires, which extended the light period, which was subsequently further amplified by lamps and candles. And over the course of the 20th century, electrical power has made it possible for people to stay up very late when it is dark\u2014working, partying, worrying\u2014to a greater extent than any other closely related species. In other words, we evolved to be diurnal, yet we are now far more nocturnal than any of our recent ancestors or close relatives. Are we adapting to nocturnality? If so, why? Does it even make any sense to speak of the human occupation of a nocturnal ape niche, despite the fact that we empirically seem to be doing just that? And if so, does it make sense to ask what the reason for it is?<\/p>\n<p class=\"import-Normal\">Third, there is a genetic phenomenon known as a selective sweep, or the hitchhiker effect. Imagine three genes\u2014A, B, and C\u2014located very closely together on a chromosome. They each have several variants, or alleles, in the population. Now, for whatever reason, it becomes beneficial to have one of the B alleles, say B4; this B4 allele is now under strong positive selection. Obviously, we will expect future generations to be characterized by mostly B4. But what was B4 attached to? Because whatever A and C alleles were adjacent to it will also be quickly spread, simply by virtue of the selection for B4. Even if the A and C alleles are not very good, they will spread because of the good B4 allele between them. Eventually the linkage groups will break up because of genetic crossing-over in future generations. But in the meantime, some random version of genes A and C are proliferating in the species simply because they are joined to superior allele B4. And clearly, the A and C alleles are there because of selection\u2014but not because of selection <em>for<\/em> them!<\/p>\n<p class=\"import-Normal\">Fourth, some features are simply consequences of other properties rather than adaptations to external conditions. We already noted the phenomenon of allometric growth, in which some physical features have to outgrow others to maintain function at an increased size. Can we ask the reason for the massive brow ridges of <em>Homo erectus<\/em>, or are brow ridges simply what you get when you have a conjunction of thick skull bones, a large face, and a sloping forehead\u2014and, thus, again would have a cause but no reason?<\/p>\n<p class=\"import-Normal\">Fifth, some features may be underutilized and on the way out. What is the reason for our two outer toes? They aren\u2019t propulsive, they don\u2019t do anything, and sometimes they\u2019re just in the way. Obviously they are there because we are descended from ancestors with five digits on their hands and feet. Is it possible that a million years from now, we will just have our three largest toes, just as the ancestors of the horse lost their digits in favor of a single hoof per limb? Or will our outer toes find another use, such as stabilizing the landings in our personal jet-packs? For the time being, we can just recognize vestigiality as another nonadaptive explanation for the presence of a given feature.<\/p>\n<p class=\"import-Normal\">Finally, Darwin himself recognized that many obvious features do not help an animal survive. Some things may instead help an animal breed. The peacock\u2019s tail feathers do not help it eat, but they do help it mate. There is competition, but only against half of the species. Darwin called this <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1084\">sexual selection<\/a><\/strong>. Its result is not a fit to the environment but, rather, a fit to the opposite sex. In some species, that is literally the case, as the male and female genitalia have specific ways of anatomically fitting together. The specific form is less important than the specific match, so inquiring about the reason for a particular form of the reproductive anatomy may be misleading. The specific form may be effectively random, as long as it fits the opposite sex and is different from the anatomies of other species. Nor is sexual selection the only form of selection that can affect the body differently from natural selection. Competition might also take place between biological units other than organisms\u2014perhaps genes, perhaps cells, or populations, or species. The spread of cultural things, such as head-binding or cheap refined fructose or forced labor, can have significant effects upon bodies, which are also not adaptations produced by natural selection. They are often adaptive physiological responses to stresses but not the products of natural selection.<\/p>\n<p class=\"import-Normal\">With so many paths available by which a physical feature might have organically arisen without having been the object of natural selection, it is unwise to assume that any individual trait is an adaptation. And that generalization applies to the best-known, best-studied, and most materially based evolutionary adaptations of our lineage. But our cultural behaviors are also highly adaptive, so what about our most familiar social behaviors? Patriarchy, hierarchy, warfare\u2014are these adaptations? Do they have reasons? Are they good for something?<\/p>\n<p class=\"import-Normal\">This is where some sloppy thinking has been troublesome. What would it mean to say that patriarchy evolved by natural selection in the human species? If, on the one hand, it means that the human mind evolved by natural selection to be able to create and survive in many different kinds of social and political regimes, of which patriarchy is one, then biological anthropologists will readily agree. If, on the other hand, it means that patriarchy evolved by natural selection, that implies that patriarchy is genetically determined (since natural selection is a genetic process) and out-reproduced the alleles for other, more egalitarian, social forms. This in turn would imply that patriarchy is an adaptation and therefore of some beneficial value in the past and has become an ingrained part of human nature today. This would be bad news, say, if you harbored ambitions of dismantling it. Dismantling patriarchy in that case would be to go against nature, a futile gesture. In other words, this latter interpretation would be a naturalistic manifesto for a conservative political platform: don\u2019t try to dismantle the patriarchy, because it is within us, the product of evolution\u2014suck it up and live with it.<\/p>\n<p class=\"import-Normal\">Here, evolution is being used as a political instrument for transforming the human genome into an imaginary glass ceiling against equality. There is thus a convergence between the pseudo-biology of crude <strong>adaptationism <\/strong>(the idea that everything is the product of natural selection) and the pseudo-biology of hereditarianism. Naturalizing inequality is not the business of evolutionary theory, and it represents a difficult moral position for a scientist to adopt, as well as a poor scientific position.<\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ffff00\"><strong style=\"font-family: 'Cormorant Garamond', serif;font-size: 1.602em;background-color: #ffff00\">Evolution of the Anthropocene (to be reviewed)<\/strong><\/span><\/p>\n<figure style=\"width: 404px\" class=\"wp-caption alignright\"><img class=\"\" src=\"https:\/\/images.squarespace-cdn.com\/content\/v1\/5915c70e59cc6830f44a9d74\/1535724432724-FJ6FIAZI1B7RR65F3FMD\/ANTH_TFOS_DAN_02_16_SRC_WEB+%282%29.jpg\" alt=\"Dandora Landfill #3, Plastics Recycling, Nairobi, Kenya 2016\" width=\"404\" height=\"302\" \/><figcaption class=\"wp-caption-text\">comes from here https:\/\/www.edwardburtynsky.com\/projects\/the-anthropocene-project<\/figcaption><\/figure>\n<p>Under the previously explored Adaptationism and Panglossian Paradigm, it is explained that human evolution is constantly occurring even throughout periods of ecological stability. While this acknowledges evolution as an ongoing process of change, it fails to explore the implications of such on the alteration of other species and ecosystems.<\/p>\n<p>The emergence of the Anthropocene, driven by human activity, though not recognized as an official epoch, is seen as a transformative event comparable to other major historical shifts such as the Ordovician Biodiversification (UNESCO, 2024). Given its scale, it is crucial to inform scholars about the impact of our social and cultural evolution on the rest of the world. Richard Robbins\u2019 Global Problems and Culture of Capitalism explains how the modern culture of consumption has been extremely successful at accommodating populations of people far larger than previously possible. Robbins claims that the globalization attributed to capitalism has allowed the world to make full use of its environmental resources, providing necessities and innovative technologies to humans all over the world (Robbins &amp; Dowty, 2019). In other words, capitalism is an anthropocentric cultural system that highly benefits humans and facilitates our survival with little regard to the development and survival of other forms of life. It would be highly relevant to introduce the idea that our cultural evolution and capacity to modify the environment to meet our needs have established new environmental conditions in which the human species' survival and reproduction rate expand at the detriment of ecosystems and endangerment of other primates and non-human species.<\/p>\n<p>According to the International Union for Conservation of Nature\u2019s Red List of Threatened Species, there are currently over 169,000 species listed, with more than 47,000 species at risk of extinction \u2014 including 41% of amphibians, 26% of mammals, 26% of freshwater fishes, 12% of birds, and many others (IUCN, 2025). Human lifestyles are causing changes that\u2014if not taken into consideration\u2014could lead to our extinction as a species. The recognition that our evolutionary behavioural development is causing environmental destruction may be the first step for our species to take accountability for the damage that it is causing to others and prevent further damage.<\/p>\n<p><span style=\"background-color: #ff99cc;font-family: 'Cormorant Garamond', serif;font-size: 1.602em;font-weight: bold\">Concluding Thoughts<\/span><\/p>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Now that you have finished reading this chapter, you are equipped to understand the historical and political dimensions of evolution. Evolution is an ongoing process of change and diversification. Evolutionary theory is a tool that we use to understand this process. The development of evolutionary theory is shaped both by scientific innovation and political engagement. Since Darwin first articulated natural selection as an observable mechanism by which species adapt to their environments, our understanding of evolution has grown. Initially, scientists focused on the adaptive aspects of evolution. However, with the emergence of genetics, our understanding of heredity and the level at which evolution acts has changed. Genetics led to a focus on the molecular dimensions of evolution. For some, this focus resulted in reductive accounts of evolution. Further developments in our understanding of evolution shifted our view to epigenetic processes and how organisms shape their own evolutionary pressures (e.g., niche construction).<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Evolutionary theory will continue to develop in the future as we invent new technologies, describe new dimensions of biology, and experience cultural changes. Current innovations in evolutionary theory are asking us to consider evolutionary forces beyond natural selection and genetics to include the ways organisms shape their environments (niche construction), inheritances beyond genetics (inclusive inheritance), constraints on evolutionary change (developmental bias), and the ability of bodies to change in response to external factors (plasticity). The future of evolutionary theory looks bright as we continue to explore these and other dimensions. Biological anthropology is well-positioned to be a lively part of this conversation, as it extends standard evolutionary theory by considering the role of culture, social learning, and human intentionality in shaping the evolutionary trajectories of humans (Zeder 2018). Remember, at root, human evolutionary theory consists of two propositions: (1) the human species is descended from other similar species and (2) natural selection has been the primary agent of biological adaptation. Pretty much everything else is subject to some degree of contestation.<\/span><\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Review Questions<\/h2>\n<ul>\n<li class=\"import-Normal\">How is the study of your ancestors biopolitical, not just biological? Does that make it less scientific or differently scientific?<\/li>\n<li class=\"import-Normal\">What was gained by reducing organisms to genotypes and species to gene pools? What is gained by reintroducing bodies and species into evolutionary studies?<\/li>\n<li class=\"import-Normal\">How do genetic or molecular studies complement anatomical studies of evolution?<\/li>\n<li class=\"import-Normal\">How are you reducible to your ancestry? If you could meet your ancestors from the year 1700 (and you would have well over a thousand of them!), would their lives be meaningfully similar to yours? Would you even be able to communicate with them?<\/li>\n<li class=\"import-Normal\">The molecular biologist Fran\u00e7ois Jacob argued that evolution is more like a tinkerer than an engineer. In what ways do we seem like precisely engineered machinery, and in what ways do we seem like jerry-rigged or improvised contraptions?<\/li>\n<li class=\"import-Normal\">How might biological anthropology contribute to future developments in evolutionary theory?<\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\">Key Terms<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\"><strong>Adaptation<\/strong>: A fit between the organism and environment.<\/p>\n<p class=\"import-Normal\"><strong>Adaptationism<\/strong>: The idea that everything is the product of natural selection.<\/p>\n<p class=\"import-Normal\"><strong>Allele<\/strong>: A genetic variant.<\/p>\n<p class=\"import-Normal\"><strong>Allometry<\/strong>: The differential growth of body parts.<\/p>\n<p class=\"import-Normal\"><strong>Canalization<\/strong>: The tendency of a growing organism to be buffered toward normal development.<\/p>\n<p class=\"import-Normal\"><strong>Epigenetics<\/strong>: The study of how genetically identical cells and organisms (with the same DNA base sequence) can nevertheless differ in stably inherited ways.<\/p>\n<p class=\"import-Normal\"><strong>Eugenics<\/strong>: An idea that was popular in the 1920s that society should be improved by breeding \u201cbetter\u201d kinds of people.<\/p>\n<p class=\"import-Normal\"><strong>Evo-devo<\/strong>: The study of the origin of form; a contraction of \u201cevolutionary developmental biology.\u201d<\/p>\n<p class=\"import-Normal\"><strong>Exaptation<\/strong>: An additional beneficial use for a biological feature.<\/p>\n<p class=\"import-Normal\"><strong>Extinction<\/strong>: The loss of a species from the face of the earth.<\/p>\n<p class=\"import-Normal\"><strong>Gene<\/strong>: A stretch of DNA with an identifiable function (sometimes broadened to include any DNA with recognizable structural features as well).<\/p>\n<p class=\"import-Normal\"><strong>Gene pool<\/strong>: Hypothetical summation of the entire genetic composition of population or species.<\/p>\n<p class=\"import-Normal\"><strong>Genotype<\/strong>: Genetic constitution of an individual organism.<\/p>\n<p class=\"import-Normal\"><strong>Hereditarianism<\/strong>: The idea that genes or ancestry is the most crucial or salient element in a human life. Generally associated with an argument for natural inequality on pseudo-genetic grounds.<\/p>\n<p class=\"import-Normal\"><strong>Hox genes<\/strong>: A group of related genes that control for the body plan of an embryo along the head-tail axis.<\/p>\n<p class=\"import-Normal\"><strong>Inheritance of acquired characteristics<\/strong>: The idea that you pass on the features that developed during your lifetime, not just your genes; also known as Lamarckian inheritance.<\/p>\n<p class=\"import-Normal\"><strong>Natural selection<\/strong>: A consistent bias in survival and fertility, leading to the overrepresentation of certain features in future generations and an improved fit between an average member of the population and the environment.<\/p>\n<p class=\"import-Normal\"><strong>Niche construction<\/strong>: The active engagement by which species transform their surroundings in favorable ways, rather than just passively inhabiting them.<\/p>\n<p class=\"import-Normal\"><strong>Phenotype<\/strong>: Observable manifestation of a genetic constitution, expressed in a particular set of circumstances. The suite of traits of an organism.<\/p>\n<p class=\"import-Normal\"><strong>Phrenology<\/strong>: The 19th-century anatomical study of bumps on the head as an indication of personality and mental abilities.<\/p>\n<p class=\"import-Normal\"><strong>Plasticity<\/strong>: The tendency of a growing organism to react developmentally to its particular conditions of life.<\/p>\n<p class=\"import-Normal\"><strong>Punctuated equilibria<\/strong>: The idea that species are stable through time and are formed very rapidly relative to their duration. (The opposite theory, that species are unstable and constantly changing through time, is called phyletic gradualism.)<\/p>\n<p class=\"import-Normal\"><strong>Scientific racism<\/strong>: The use of pseudoscientific evidence to support or legitimize racial hierarchy and inequality.<\/p>\n<p class=\"import-Normal\"><strong>Sexual selection<\/strong>: Natural selection arising through preference by one sex for certain characteristics in individuals of the other sex.<\/p>\n<p class=\"import-Normal\"><strong>Species selection<\/strong>: A postulated evolutionary process in which selection acts on an entire species population, rather than individuals.<\/p>\n<h2 class=\"import-Normal\">About the Authors<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image2-4-1.jpg\" alt=\"A bearded man wearing glasses smiles at the camera. \" width=\"202\" height=\"218\" \/><\/p>\n<h3 class=\"import-Normal\"><strong>Jonathan Marks, Ph.D.<\/strong><\/h3>\n<p class=\"import-Normal\">University of North Carolina at Charlotte, <a class=\"rId41\" href=\"mailto:jmarks@uncc.edu\">jmarks@uncc.edu<\/a><\/p>\n<p class=\"import-Normal\">Jonathan Marks is Professor of Anthropology at the University of North Carolina at Charlotte. He has published many books and articles on broad aspects of biological anthropology. In 2006 he was elected a Fellow of the American Association for the Advancement of Science. In 2012 he was awarded the First Citizen\u2019s Bank Scholar\u2019s Medal from UNC Charlotte. In recent years he has been a Visiting Research Fellow at the ESRC Genomics Forum in Edinburgh, a Visiting Research Fellow at the Max Planck Institute for the History of Science in Berlin, and a Templeton Fellow at the Institute for Advanced Study at Notre Dame. His work has received the W. W. Howells Book Prize and the General Anthropology Division Prize for Exemplary Cross-Field Scholarship from the American Anthropological Association as well as the J. I. Staley Prize from the School for Advanced Research. Two of his books are titled <em>What It Means to Be 98% Chimpanzee<\/em> and <em>Why I Am Not a Scientist<\/em>, but actually he is about 98 percent scientist and not a chimpanzee.<\/p>\n<p class=\"import-Normal\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image10-3.jpg\" alt=\"A bearded man wearing a fedora hat looks off in the distance. \" width=\"232\" height=\"232\" \/><\/p>\n<h3 class=\"import-Normal\"><strong>Adam P. Johnson, M.A.<\/strong><\/h3>\n<p class=\"import-Normal\">University of North Carolina at Charlotte\/University of Texas at San Antonio, <a class=\"rId43\" href=\"mailto:ajohn344@uncc.edu\">ajohn344@uncc.edu<\/a><\/p>\n<p class=\"import-Normal\">Adam Johnson is a doctoral candidate at the University of Texas at San Antonio and part-time lecturer at the University of North Carolina at Charlotte. He earned his M.A. in anthropology at UNC-Charlotte in 2017 and will complete his Ph.D. in anthropology at UTSA by 2024. His interests include human-animal relations, science studies, primate behavior, ecology, and the history of anthropology. His recent research project analyzes the social, historical, political, and evolutionary dimensions that shape human-javelina encounters. His goal is to understand how humans and animals find ways to get along in a precarious world.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration <strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\">Ackermann, Rebecca Rogers, Alex Mackay, and Michael L. Arnold. 2016. \u201cThe Hybrid Origin of \u2018Modern\u2019 Humans.\u201d <em>Evolutionary Biology<\/em> 43 (1): 1\u201311.<\/p>\n<p class=\"import-Normal\">Bateson, Patrick, and Peter Gluckman. 2011. <em>Plasticity, Robustness, Development and Evolution<\/em>. New York: Cambridge University Press.<\/p>\n<p class=\"import-Normal\">Cosans, Christopher E. 2009. <em>Owen's Ape and Darwin's Bulldog: Beyond Darwinism and Creationism<\/em>. Bloomington, IN: Indiana University Press.<\/p>\n<p class=\"import-Normal\">Desmond, Adrian, and James Moore. 2009. <em>Darwin's Sacred Cause: How a Hatred of Slavery Shaped Darwin's Views on Human Evolution<\/em>. New York: Houghton Mifflin Harcourt.<\/p>\n<p class=\"import-Normal\">Dobzhansky, Theodosius, Francisco J. Ayala, G. Ledyard Stebbins, and James W. Valentine. 1977. <em>Evolution<\/em>. San Francisco: W.H. Freeman and Company.<\/p>\n<p class=\"import-Normal\">Fuentes, Agust\u00edn. 2017. <em>The Creative Spark: How Imagination Made Humans Exceptional<\/em>. New York: Dutton.<\/p>\n<p class=\"import-Normal\">Gould, Stephen J. 2003.<em> The Structure of Evolutionary Theory<\/em>. Cambridge, MA: Harvard University Press.<\/p>\n<p class=\"import-Normal\">Haraway, Donna J. 1989. <em>Primate Visions: Gender, Race, and Nature in the World of Modern Science<\/em>. New York: Routledge.<\/p>\n<p class=\"import-Normal\">Huxley, Thomas. 1863. <em>Evidence as to Man's Place in Nature<\/em>. London: Williams &amp; Norgate.<\/p>\n<p class=\"import-Normal\">Jablonka, Eva, and Marion J. Lamb. 2005. <em>Evolution in Four Dimensions: Genetic, Epigenetic, Behavioral, and Symbolic Variation in the History of Life<\/em>. Cambridge, MA: The MIT Press.<\/p>\n<p class=\"import-Normal\">Kuklick, Henrika, ed. 2008. <em>A New History of Anthropology<\/em>. New York: Blackwell.<\/p>\n<p class=\"import-Normal\">Laland, Kevin N., Tobias Uller, Marcus W. Feldman, Kim Sterelny, Gerd B. Muller, Armin Moczek, Eva Jablonka, and John Odling-Smee. 2015. \u201cThe Extended Evolutionary Synthesis: Its Structure, Assumptions and Predictions.\u201d <em>Proceedings of the Royal Society, Series B<\/em> 282 (1813): 20151019.<\/p>\n<p class=\"import-Normal\">Lamarck, Jean Baptiste. 1809. <em>Philosophie Zoologique<\/em>. Paris: Dentu.<\/p>\n<p class=\"import-Normal\">Landau, Misia. 1991. <em>Narratives of Human Evolution<\/em>. New Haven: Yale University Press.<\/p>\n<p class=\"import-Normal\">Lee, Sang-Hee. 2017. <em>Close Encounters with Humankind: A Paleoanthropologist Investigates Our Evolving Species<\/em>. New York: W. W. Norton.<\/p>\n<p class=\"import-Normal\">Livingstone, David N. 2008. <em>Adam's Ancestors: Race, Religion, and the Politics of Human Origins<\/em>. Baltimore: Johns Hopkins University Press.<\/p>\n<p class=\"import-Normal\">Marks, Jonathan. 2015. <em>Tales of the Ex-Apes: How We Think about Human Evolution<\/em>. Berkeley, CA: University of California Press.<\/p>\n<p class=\"import-Normal\">Pigliucci, Massimo. 2009. \u201cThe Year in Evolutionary Biology 2009: An Extended Synthesis for Evolutionary Biology.\u201d <em>Annals of the New York Academy of Sciences<\/em> 1168: 218\u2013228.<\/p>\n<p class=\"import-Normal\">Simpson, George Gaylord. 1949. <em>The Meaning of Evolution: A Study of the History of Life and of Its Significance for Man<\/em>. New Haven: Yale University Press.<\/p>\n<p class=\"import-Normal\">Sommer, Marianne. 2016.<em> History Within: The Science, Culture, and Politics of Bones, Organisms, and Molecules<\/em>. Chicago: University of Chicago Press.<\/p>\n<p class=\"import-Normal\">Stoczkowski, Wiktor. 2002. <em>Explaining Human Origins: Myth, Imagination and Conjecture<\/em>. New York: Cambridge University Press.<\/p>\n<p class=\"import-Normal\">Tattersall, Ian, and Rob DeSalle. 2019. <em>The Accidental Homo sapiens: Genetics, Behavior, and Free Will<\/em>. New York: Pegasus.<\/p>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\">Barton, Robert A. 1996. \"Neocortex Size and Behavioural Ecology in Primates.\" <em>Proceedings of the Royal Society of London. Series B: Biological Sciences<\/em> 263 (1367): 173\u2013177.<\/p>\n<p class=\"import-Normal\">Bodmer, Walter, and Robin McKie. 1997. <em>The Book of Man: The Hman Genome Project and the Quest to Discover our Genetic Heritage.<\/em> Oxford University Press.<\/p>\n<p class=\"import-Normal\">Darwin, Charles. 1859.<em> On the Origin of Species by Means of Natural Selection, or, the Preservation of Favoured Races in the Struggle for Life<\/em>. London: J. Murray.<\/p>\n<p class=\"import-Normal\">Darwin, Charles. 1871. <em>The Descent of Man, and Selection in Relation to Sex.<\/em> London: J. Murray.<\/p>\n<p class=\"import-Normal\">Dawkins, Richard. 1976. <em>The Selfish Gene. <\/em>Oxford University Press.<\/p>\n<p class=\"import-Normal\">Deacon, T. W. 1998. <em>The Symbolic Species: The Co-evolution of Language and the Brain<\/em>. W. W. Norton &amp; Company.<\/p>\n<p class=\"import-Normal\">Eldredge, N., and S. J. Gould. 1972. \"Punctuated Equilibria: An Alternative to Phyletic Gradualism.\" In <em>Models in Paleobiology<\/em>, edited by T. J. Schopf, 82\u2013115. San Francisco: W. H. Freeman.<\/p>\n<p class=\"import-Normal\">Gould, Stephen J. 2003.<em> The Structure of Evolutionary Theory<\/em>. Cambridge, MA: Harvard University Press.<\/p>\n<p class=\"import-Normal\">Gould, Stephen J. 1996. <em>Mismeasure of Man<\/em>. New York: WW Norton &amp; Company.<\/p>\n<p class=\"import-Normal\">Gould, Stephen Jay, and Richard C. Lewontin. 1979. \"The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme.\" <em>Proceedings of the Royal Society of London. Series B: Biological Sciences<\/em> 205 (1151): 581\u2013598.<\/p>\n<p class=\"import-Normal\">Haeckel, Ernst. 1868. <em>Nat\u00fcrliche Sch\u00f6pfungsgeschichte<\/em>. Berlin: Reimer.<\/p>\n<p class=\"import-Normal\">Huxley, Thomas Henry. 1863. <em>Evidence as to Man\u2019s Place in Nature. <\/em>London: Williams and Norgate.<\/p>\n<p class=\"import-Normal\">Kaufman, Thomas C., Mark A. Seeger, and Gary Olsen. 1990. \"Molecular and Genetic Organization of the Antennapedia Gene Complex of <em>Drosophila melanogaster<\/em>.\" <em>Advances in Genetics<\/em> 27: 309\u2013362.<\/p>\n<p class=\"import-Normal\">Kellogg, Vernon. 1917. <em>Headquarters Nights<\/em>. Boston: The Atlantic Monthly Press.<\/p>\n<p class=\"import-Normal\">Kevles, Daniel J., and Leroy Hood. 1993. <em>The Code of Codes: Scientific and Social Issues in the Human Genome Project<\/em>. Cambridge, MA: Harvard University Press.<\/p>\n<p class=\"import-Normal\">Lewontin, Richard, Steven Rose, and Leon Kamin. 2017. <em>Not in Our Genes\u202f: Biology, Ideology, and Human Nature<\/em>, 2nd ed. Chicago: Haymarket Books.<\/p>\n<p class=\"import-Normal\">Lloyd, Elisabeth A., and Stephen J. Gould. 1993. \"Species Selection on Variability.\" <em>Proceedings of the National Academy of Sciences<\/em> 90 (2): 595\u2013599.<\/p>\n<p class=\"import-Normal\">Marks, Jonathan. 2015. \u201cThe Biological Myth of Human Evolution.\u201d In <em>Biologising the Social Sciences: Challenging Darwinian and Neuroscience Explanations<\/em>, edited by David Canter and David A. Turner, 59\u201378. London: Routledge.<\/p>\n<p class=\"import-Normal\">Monypenny, William Flavelle, and George Earle Buckle. 1929. <em>The Life of Benjamin Disraeli, Earl of Beaconsfield, Volume II: 1860\u20131881<\/em>. London: John Murray.<\/p>\n<p class=\"import-Normal\">Potts, Rick. 1998. \u201cVariability Selection in Hominid Evolution.\u201d <em>Evolutionary Anthropology <\/em><em>7<\/em><em>:<\/em> 81\u201396.<\/p>\n<p class=\"import-Normal\">Punnett, R. C. 1905. <em>Mendelism<\/em>. Cambridge: Macmillan and Bowes.<\/p>\n<p class=\"import-Normal\">Shapiro, Robert. 1991. <em>The Human Blueprint: The Race to Unlock the Secrets of Our Genetic Script.<\/em> New York: St. Martin\u2019s Press.<\/p>\n<p class=\"import-Normal\">Shultz, Susanne, Emma Nelson, and Robin Dunbar. 2012. \"Hominin Cognitive Evolution: Identifying Patterns and Processes in the Fossil and Archaeological Record.\" <em>Philosophical Transactions of the Royal Society B: Biological Sciences<\/em> 367 (1599): 2130\u20132140.<\/p>\n<p class=\"import-Normal\">Spencer, Herbert. 1864. <em>Principles of Biology.<\/em> London: Williams and Norgate.<\/p>\n<p class=\"import-Normal\">Watson, James D. 1990. \"The Human Genome Project: Past, Present, and Future.\" <em>Science<\/em> 248 (4951): 44\u201349.<\/p>\n<p class=\"import-Normal\">Yengo, L., Vedantam, S., Marouli, E., Sidorenko, J., Bartell, E., Sakaue, S., Graff, M., Eliasen, A.U., Jiang, Y., Raghavan, S. and Miao, J., 2022. A saturated map of common genetic variants associated with human height. <em>Nature<\/em>, <em>610 <\/em>(7933): 704-712.<\/p>\n<p class=\"import-Normal\">Zeder, Melinda A. 2018. \"Why Evolutionary Biology Needs Anthropology: Evaluating Core Assumptions of the Extended Evolutionary Synthesis.\" <em>Evolutionary Anthropology: Issues, News, and Reviews<\/em> 27 (6): 267\u2013284.<\/p>\n<\/div>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1769\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1769\"><div tabindex=\"-1\"><p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kerryn Warren, Ph.D., Grad Coach International<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lindsay Hunter, M.A., University of Iowa<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Navashni Naidoo, M.Sc., University of Cape Town<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Silindokuhle Mavuso, M.Sc., University of Witwatersrand<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\"><em>This chapter is a revision from <\/em><em>\"<\/em><a class=\"rId7\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\"><em>Chapter 9: Early Hominins<\/em><\/a><em>\" <\/em><em>by Kerryn Warren, K. Lindsay Hunter, Navashni Naidoo, Silindokuhle Mavuso, Kimberleigh Tommy, Rosa Moll, and Nomawethu Hlazo<\/em><em>. In <\/em><a class=\"rId8\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, first edition<\/em><\/a><em>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under <\/em><a class=\"rId9\" style=\"color: #000000\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\"><em>CC BY-NC 4.0<\/em><\/a><em>. <\/em><\/span><\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\"><span style=\"color: #000000\"><span style=\"color: #ffffff\">Learning Objectives<\/span><br \/>\n<\/span><\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li><span style=\"color: #000000\">Understand what is meant by \u201cderived\u201d and \u201cancestral\u201d traits and why this is relevant for understanding early hominin evolution.<\/span><\/li>\n<li><span style=\"color: #000000\">Understand changing paleoclimates and paleoenvironments as potential factors influencing early hominin adaptations.<\/span><\/li>\n<li><span style=\"color: #000000\">Describe the anatomical changes associated with bipedalism and dentition in early hominins, as well as their implications..<\/span><\/li>\n<li><span style=\"color: #000000\">Describe early hominin genera and species, including their currently understood dates and geographic expanses.<\/span><\/li>\n<li><span style=\"color: #000000\">Describe the earliest stone tool techno-complexes and their impact on the transition from early hominins to our genus.<\/span><\/li>\n<\/ul>\n<\/div>\n<\/div>\n<h2 class=\"__UNKNOWN__\"><span style=\"color: #000000\">Defining Hominins<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">It is through our study of our hominin ancestors and relatives that we are exposed to a world of \u201cmight have beens\u201d: of other paths not taken by our species, other ways of being human. But to better understand these different evolutionary trajectories, we must first define the terms we are using. If an imaginary line were drawn between ourselves and our closest relatives, the great apes, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1200\">bipedalism<\/a><\/strong> (or habitually walking upright on two feet) is where that line would be. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1270\">Hominin<\/a><\/strong>, then, means everyone on \u201cour\u201d side of the line: humans and all of our extinct bipedal ancestors and relatives since our divergence from the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1272\">last common ancestor (LCA)<\/a><\/strong> we share with chimpanzees.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Historic interpretations of our evolution, prior to our finding of early hominin <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1274\"><strong>fossils<\/strong><\/a>, varied. Debates in the mid-1800s regarding hominin origins focused on two key issues:<\/span><\/p>\n<ul>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">Where did we evolve?<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">Which traits evolved first?<\/span><\/li>\n<\/ul>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff00ff\">Charles Darwin hypothesized that we evolved in Africa, as he was convinced that we shared greater commonality with chimpanzees and gorillas on the continent (Darwin 1871). Others, such as Ernst Haeckel and Eug\u00e8ne Dubois, insisted that we were closer in affinity to orangutans and that we evolved in Eurasia where, until the discovery of the Taung Child in South Africa in 1924, all humanlike fossils (of Neanderthals and <em>Homo erectus<\/em>) had been found (Shipman 2002).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Within this conversation, naturalists and early <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1276\">paleoanthropologists<\/a><\/strong> (people who study human evolution) speculated about which human traits came first. These included the evolution of a big brain (<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1278\"><strong>encephalization<\/strong><\/a>), the evolution of the way in which we move about on two legs (bipedalism), and the evolution of our flat faces and small teeth (indications of dietary change). Original hypotheses suggested that, in order to be motivated to change diet and move about in a bipedal fashion, the large brain needed to have evolved first, as is seen in the fossil species mentioned above.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">However, we now know that bipedal locomotion is one of the first things that evolved in our lineage, with early relatives having more apelike dentition and small brain sizes. While brain size expansion is seen primarily in our genus, <em>Homo<\/em>, earlier hominin brain sizes were highly variable between and within taxa, from 300 cc (cranial capacity, cm<sup>3<\/sup>), estimated in <em>Ardipithecus<\/em>, to 550 cc, estimated in <em>Paranthropus boisei<\/em>. The lower estimates are well within the range of variation of nonhuman extant great apes. In addition, body size variability also plays a role in the interpretation of whether brain size could be considered large or small for a particular species or specimen. In this chapter, we will tease out the details of early hominin evolution in terms of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1280\"><strong>morphology<\/strong><\/a> (i.e. the study of the form, size, or shape of things; in this case, skeletal parts).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">We also know that early human evolution occurred in a very complicated fashion. There were multiple species (multiple genera) that featured diversity in their diets and locomotion. Specimens have been found all along the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1282\"><strong>East African Rift System <\/strong>(<strong>EARS)<\/strong><\/a>; that is, in Ethiopia, Kenya, Tanzania, and Malawi; see Figure 9.1), in limestone caves in South Africa, and in Chad. Dates of these early relatives range from around 7 million years ago (mya) to around 1 mya, overlapping temporally with members of our genus, <em>Homo<\/em>.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 610px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/06\/image38.png\" alt=\"Patchy green mountain alongside a deep sandy valley in East Africa.\" width=\"610\" height=\"277\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.1: East African Rift System (EARS). Credit: <a href=\"https:\/\/www.flickr.com\/photos\/ninara\/8624605781\/in\/photolist-x2yH7-x2yHe-VfVWuD-e98mPF-SzzjsU-2bsBZhC-2hHec7m-xtJ7Ez-NXnXvh-7Yg3uo-2cS3FgG-2hjo1Dc-2hjGoTS-nnumi8-82U66W-dMNn7B-8jdVbd-NWDg8-NW6fj-ebhx5w-bkFv1G-Ct5ZD-5JQk8A-y6TgAc-x9k6oe-2ebLTDC-WcPMnJ-2ekh6CS-Cu3LH-xNHDFK-9RUsZi-94jVt4-P46uiB-QFyjyE-crU8N7-5JLJKV-2ekSgk8-5JL454-2cPgZrF-2bHfQZu-dMTVPN-6yUbeN-jzMicQ-48XjU9-2etR2Ze-Styrvw-crU7V7-2wakq3-crU6Z1-2etR2XR\/\">IMG_1696 Great Rift Valley<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/ninara\/\">Ninara<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\">CC BY 2.0 License<\/a>.<\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Yet there is still so much to understand. Modern debates now look at the relatedness of these species to us and to one another, and they consider which of these species were able to make and use tools. As a result, every <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1284\">site<\/a><\/strong> discovery in the patchy hominin fossil record tells us more about our evolution. In addition, recent scientific techniques (not available even ten years ago) provide new insights into the diets, environments, and lifestyles of these ancient relatives.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In the past, <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1288\"><strong>taxonom<\/strong><strong>y<\/strong><\/a> was primarily based on morphology. Today it is tied to known relationships based on molecular <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1290\"><strong>phylogeny<\/strong><\/a> (e.g., based on DNA) or a combination of the two. This is complicated when applied to living <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1286\"><strong>taxa<\/strong><\/a>, but becomes much more difficult when we try to categorize ancestor-descendant relationships for long-extinct species whose molecular information is no longer preserved. We therefore find ourselves falling back on morphological comparisons, often of teeth and partially fossilized skeletal material.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">It is here that we turn to the related concepts of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1292\">cladistics<\/a> <\/strong>and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1294\">phylogenetics<\/a><\/strong><strong>. <\/strong><span style=\"background-color: #ff99cc\">Cladistics groups organisms according to their last common ancestors based on shared <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1126\">derived traits<\/a><\/strong>. <\/span>In the case of early hominins, these are often morphological traits that differ from those seen in earlier populations. These new or modified traits provide evidence of evolutionary relationships, and organisms with the same derived traits are grouped in the same <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1116\">clade<\/a> <\/strong>(Figure 9.2). <span style=\"background-color: #ff99cc\">For example, if we use feathers as a trait, we can group pigeons and ostriches into the clade of birds.<\/span> In this chapter, we will examine the grouping of the Robust Australopithecines, whose cranial and dental features differ from those of earlier hominins, and therefore are considered derived.<\/span><\/p>\n<figure style=\"width: 708px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image48.png\" alt=\"Phylogenetic tree shows clades and non clade groupings.\" width=\"708\" height=\"192\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.2: Clades refer to groups of species or taxa that share a common ancestor. In <span class=\"ILfuVd\" lang=\"en\"><span class=\"hgKElc\">a phylogeny, a clade is a complete group of lineages, including their last common ancestor. Groupings that do not include a common ancestor and <em>all<\/em> of its descendants are not clades. <\/span><\/span>Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\">Clades (Figure 9.2)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Katie Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Dig Deeper: Problems Defining Hominin Species<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000\">It is worth noting that species designations for early hominin specimens are often highly contested. This is due to the fragmentary nature of the fossil record, the large timescale (millions of years) with which paleoanthropologists need to work, and the difficulty in evaluating whether morphological differences and similarities are due to meaningful phylogenetic or biological differences or subtle differences\/variation in niche occupation or time. In other words, do morphological differences really indicate different species? How would classifying species in the paleoanthropological record compare with classifying living species today, for whom we can sequence genomes and observe lifestyles?<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">There are also broader philosophical differences among researchers when it comes to paleo-species designations. Some scientists, known as \u201c<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1296\"><strong>lumpers<\/strong>,<\/a>\u201d argue that large variability is expected among multiple populations in a given species over time. These researchers will therefore prefer to \u201clump\u201d specimens of subtle differences into single taxa. Others, known as \u201c<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1298\"><strong>splitters<\/strong><\/a>,\u201d argue that species variability can be measured and that even subtle differences can imply differences in niche occupation that are extreme enough to mirror modern species differences. In general, splitters would consider geographic differences among populations as meaning that a species is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1300\">polytypic<\/a><\/strong> <span style=\"background-color: #ff99cc\">(i.e., capable of interacting and breeding biologically but having morphological population differences)<\/span>. This is worth keeping in mind when learning about why species designations may be contested.<\/span><\/p>\n<figure style=\"width: 520px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image8.jpg\" alt=\"A graph shows a curved line depicting changes in morphology among two species over time.\" width=\"520\" height=\"292\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.3: This graph demonstrates the concept of a chronospecies, where one species (Species A) \u201cevolves\u201d into another (Species B). Credit: Chronospecies original to Explorations: An Open Invitation to Biological Anthropology, 2nd edition by Kerryn Warren is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\"><span style=\"color: #000000\">This further plays a role in evaluating ancestry. Debates over which species \u201cgave rise\u201d to which continue to this day. It is common to try to create \u201clineages\u201d of species to determine when one species evolved into another over time. We refer to these as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1302\"><strong>chronospecies<\/strong><\/a> (Figure 9.3). Constructed hominin phylogenetic trees are routinely variable, changing with new specimen discoveries, new techniques for evaluating and comparing species, and, some have argued, nationalist or biased interpretations of the record. More recently, some researchers have shifted away from \u201ctreelike\u201d models of ancestry toward more nuanced metaphors such as the \u201cbraided stream,\u201d where some levels of interbreeding among species and populations are seen as natural processes of evolution.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Finally, it is worth considering the process of fossil discovery and publication. Some fossils are easily diagnostic to a species level and allow for easy and accurate interpretation. Some, however, are more controversial. This could be because they do not easily preserve or are incomplete, making it difficult to compare and place within a specific species (e.g., a fossil of a patella or knee bone). Researchers often need to make several important claims when announcing or publishing a find: a secure date (if possible), clear association with other finds, and an adequate comparison among multiple species (both extant and fossil). Therefore, it is not uncommon that an important find was made years before it is scientifically published.<\/span><\/p>\n<\/div>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Paleoenvironment and Hominin Evolution<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There is no doubt that one of the major selective pressures in hominin evolution is the environment. <span style=\"background-color: #ff9900\"><span style=\"background-color: #ffff00\">Large-scale changes in global and regional climate, as well as alterations to the environment, are<\/span><span style=\"background-color: #ffff00\"> (<\/span><\/span><em style=\"background-color: transparent\"><span style=\"text-decoration: underline\">thought to be<\/span>)<\/em><span style=\"background-color: #ffff00\"><del> all<\/del> linked to (all) hominin diversification, dispersal, and extinction<\/span> (Maslin et al. 2014). Environmental reconstructions often use modern analogues. Let us take, for instance, the hippopotamus. It is an animal that thrives in environments that have abundant water to keep its skin cool and moist. If the environment for some reason becomes drier, it is expected that hippopotamus populations will reduce. If a drier environment becomes wetter, it is possible that hippopotamus populations may be attracted to the new environment and thrive. Such instances have occurred multiple times in the past, and the bones of some <strong style=\"background-color: transparent\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1308\">fauna<\/a><\/strong> (i.e., animals, like the hippopotamus) that are sensitive to these changes give us insights into these events.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Yet reconstructing a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1304\">paleoenvironment<\/a><\/strong> relies on a range of techniques, which vary depending on whether research interests focus on local changes or more global environmental changes\/reconstructions. For local environments (such as a single site or region), comparing the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1306\">faunal assemblages<\/a> <\/strong>(collections of fossils of animals found at a site) with animals found in certain modern environments allows us to determine if past environments mirror current ones in the region. Changes in the faunal assemblages, as well as when they occur and how they occur, tell us about past environmental changes. Other techniques are also useful in this regard. Chemical analyses, for instance, can reveal the diets of individual fauna, providing clues as to the relative wetness or dryness of their environment (e.g., nitrogen <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1310\"><strong>isotopes<\/strong><\/a>; Kingston and Harrison 2007).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Global climatic changes in the distant past, which fluctuated between being colder and drier and warmer and wetter on average, would have global implications for environmental change (Figure 9.4). These can be studied by comparing marine core and terrestrial soil data across multiple sites. These techniques are based on chemical analysis, such as examination of the nitrogen and oxygen isotopes in shells and sediments. Similarly, analyzing pollen grains shows which kinds of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1312\"><strong>flora<\/strong> <\/a> survived in an environment at a specific time period. There are multiple lines of evidence that allow us to visualize global climate trends over millions of years (although it should be noted that the direction and extent of these changes could differ by geographic region).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 649px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image12-1-1.png\" alt=\"Chart shows cyclical carbon dioxide levels from 800,000 years ago until today.\" width=\"649\" height=\"406\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.4: This graph, based on the comparison of atmospheric samples contained in ice cores and more recent direct measurements, illustrates how atmospheric CO\u2082 has fluctuated over time and increased sharply since the Industrial Revolution. The graph also shows that since 800,000ya (and before) atmospheric CO\u2082 has never exceeded 300 parts per million (ppm). In 1950 it was 310ppm. Today atmospheric CO\u2082 has spiked to over 410 ppm. Credit: <a href=\"https:\/\/climate.nasa.gov\/evidence\/\">CO\u2082 increase since the Industrial Revolution<\/a> by <a href=\"https:\/\/www.nasa.gov\/\">NASA<\/a> is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a> and is used within <a href=\"https:\/\/www.nasa.gov\/multimedia\/guidelines\/index.html\">NASA guidelines on re-use<\/a>. Original from Luthi, D., et al.. 2008; Etheridge, D.M., et al. 2010; Vostok ice core data\/J.R. Petit et al.; NOAA Mauna Loa CO<a href=\"https:\/\/climate.nasa.gov\/evidence\/\">\u2082<\/a> record..<\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Both local and global climatic\/environmental changes have been used to understand factors affecting our evolution (DeHeinzelin et al. 1999; Kingston 2007). Environmental change acts as an important factor regarding the onset of several important hominin traits seen in early hominins and discussed in this chapter. Namely, the environment has been interpreted as the following:<\/span><\/p>\n<ul>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">the driving force behind the evolution of bipedalism,<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">the reason for change and variation in early hominin diets, and<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">the diversification of multiple early hominin species.<\/span><\/li>\n<\/ul>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are numerous hypotheses regarding how climate has driven and continues to drive human evolution. Here, we will focus on just three popular hypotheses.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Savannah Hypothesis (or Aridity Hypothesis)<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The hypothesis:<\/strong> This popular theory suggests that the expansion of the savannah (or less densely forested, drier environments) forced early hominins from an <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1314\"><strong>arboreal<\/strong> <\/a> lifestyle (one living in trees) to a terrestrial one where bipedalism was a more efficient form of locomotion (Figure 9.5). It was first proposed by Darwin (1871) and supported by anthropologists like Raymond Dart (1925). However, this idea was supported by little fossil or paleoenvironmental evidence and was later refined as the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1318\">Aridity Hypothesis<\/a><\/strong>. This hypothesis states that the long-term <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1316\">aridification<\/a><\/strong> and, thereby, expansion of savannah biomes were drivers in diversification in early hominin evolution (deMenocal 2004; deMenocal and Bloemendal 1995). It advocates for periods of accelerated aridification leading to early hominin speciation events.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 647px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image46.png\" alt=\"Photograph showing a dry, open savannah environment.\" width=\"647\" height=\"486\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.5: The African savannah grew during early hominin evolution. This may have forced early hominins from an arboreal lifestyle to a terrestrial one, where bipedalism was a more efficient form of locomotion. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:African_savannah_@_Masai_Mara_(21308330314).jpg\">African savannah @ Masai Mara (21308330314)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/132394214@N04\">Leo Li<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\">CC BY 2.0 License<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The evidence:<\/strong> While early bipedal hominins are often associated with wetter, more closed environments (i.e., not the Savannah Hypothesis), both marine and terrestrial records seem to support general cooling, drying conditions, with isotopic records indicating an increase in grasslands (i.e., colder and wetter climatic conditions) between 8 mya and 6 mya across the African continent (Cerling et al. 2011). This can be contrasted with later climatic changes derived from aeolian dust records (sediments transported to the site of interest by wind), which demonstrate increases in seasonal rainfall between 3 mya and 2.6 mya, 1.8 mya and 1.6 mya, and 1.2 mya and 0.8 mya (deMenocal 2004; deMenocal and Bloemendal 1995).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Interpretation(s):<\/strong> Despite a relatively scarce early hominin record, it is clear that two important factors occur around the time period in which we see increasing aridity. The first factor is the diversification of taxa, where high morphological variation between specimens has led to the naming of multiple hominin genera and species. The second factor is the observation that the earliest hominin fossils appear to have traits associated with bipedalism and are dated to around the drying period (as based on isotopic records). Some have argued that it is more accurately a combination of bipedalism and arboreal locomotion, which will be discussed later. However, the local environments in which these early specimens are found (as based on the faunal assemblages) do not appear to have been dry.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Turnover Pulse Hypothesis<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The hypothesis:<\/strong> In 1985, paleontologist Elisabeth Vbra noticed that in periods of extreme and rapid climate change, <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1326\"><strong>ungulates<\/strong><\/a> (hoofed mammals of various kinds) that had generalized diets fared better than those with specialized diets (Vrba 1988, 1998). <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1322\"><strong>Specialist<\/strong><\/a> eaters <span style=\"background-color: #ff99cc\">(those who rely primarily on specific food types)<\/span> faced extinction at greater rates than their <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1320\">generalist<\/a> <\/strong><span style=\"background-color: #ff99cc\">(those who can eat more varied and variable diets) <\/span>counterparts because they were unable to adapt to new environments (Vrba 2000). Thus, periods with extreme climate change would be associated with high <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1324\"><strong>faunal turnover<\/strong><\/a>: that is, the extinction of many species and the speciation, diversification, and migration of many others to occupy various niches.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The evidence:<\/strong> The onset of the<strong> <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1400\">Quaternary Ice Age<\/a><\/strong>, between 2.5 mya and 3 mya, brought extreme global, cyclical <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1328\"><strong>interglacial<\/strong> <\/a> and <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1330\"><strong>glacial<\/strong><\/a> periods (warmer, wetter periods with less ice at the poles, and colder, drier periods with more ice near the poles). Faunal evidence from the Turkana basin in East Africa indicates multiple instances of faunal turnover and extinction events, in which global climatic change resulted in changes from closed\/forested to open\/grassier habitats at single sites (Behrensmeyer et al. 1997; Bobe and Behrensmeyer 2004). Similarly, work in the Cape Floristic Belt of South Africa shows that extreme changes in climate play a role in extinction and migration in ungulates. While this theory was originally developed for ungulates, its proponents have argued that it can be applied to hominins as well. However, the link between climate and speciation is only vaguely understood (Faith and Behrensmeyer 2013).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Interpretation(s):<\/strong> While the evidence of rapid faunal turnover among ungulates during this time period appears clear, there is still some debate around its usefulness as applied to the paleoanthropological record. Specialist hominin species do appear to exist for long periods of time during this time period, yet it is also true that <em>Homo<\/em>, a generalist genus with a varied and adaptable diet, ultimately survives the majority of these fluctuations, and the specialists appear to go extinct.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Variability Selection Hypothesis<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The hypothesis: <\/strong>This hypothesis was first articulated by paleoanthropologist Richard Potts (1998). It links the high amount of climatic variability over the last 7 million years to both behavioral and morphological changes. Unlike previous notions, this hypothesis states that hominin evolution does not respond to habitat-specific changes or to specific aridity or moisture trends. Instead, long-term environmental unpredictability over time and space influenced morphological and behavioral adaptations that would help hominins survive, regardless of environmental context (Potts 1998, 2013). The Variability Selection Hypothesis states that hominin groups would experience varying degrees of natural selection due to continually changing environments and potential group isolation. This would allow certain groups to develop genetic combinations that would increase their ability to survive in shifting environments. These populations would then have a genetic advantage over others that were forced into habitat-specific adaptations (Potts 2013).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The evidence:<\/strong> The evidence for this theory is similar to that for the Turnover Pulse Hypothesis: large climatic variability and higher survivability of generalists versus specialists. However, this hypothesis accommodates for larger time-scales of extinction and survival events.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Interpretation(s):<\/strong> In this way, the Variability Selection Hypothesis allows for a more flexible interpretation of the evolution of bipedalism in hominins and a more fluid interpretation of the Turnover Pulse Hypothesis, where species turnover is meant to be more rapid. In some ways, this hypothesis accommodates both environmental data and our interpretations of an evolution toward greater variability among species and the survivability of generalists.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000;background-color: #ff99cc\"><strong>Paleoenvironment Summary<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff99cc\">Some hypotheses presented in this section pay specific attention to habitat (Savannah Hypothesis) while others point to large-scale climatic forces (Variability Selection Hypothesis). Some may be interpreted to describe the evolution of traits such as bipedalism (Savannah Hypothesis), and others generally explain the diversification of early hominins (Turnover Pulse and Variability Selection Hypotheses). While there is no consensus as to how the environment drove our evolution, it is clear that the environment shaped both habitat and resource availability in ways that would have influenced our early ancestors physically and behaviorally.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Derived Adaptations: Bipedalism<br \/>\n<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The unique form of locomotion exhibited by modern humans, called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1332\">obligate bipedalism<\/a><\/strong>, is important in distinguishing our species from the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1334\"><strong>extant<\/strong><\/a> (living) great apes. The ability to walk habitually upright is thus considered one of the defining attributes of the hominin lineage. We also differ from other animals that walk bipedally (such as kangaroos) in that we do not have a tail to balance us as we move.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The origin of bipedalism in hominins has been debated in paleoanthropology, but at present there are two main <span style=\"background-color: #ff99cc\">ideas<\/span>: <span style=\"text-decoration: underline\">(<em>theories<\/em>)<\/span><\/span><\/p>\n<ol>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">early hominins initially lived in trees, but increasingly started living on the ground, so we were a product of an arboreal last common ancestor (LCA) or,<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">our LCA was a terrestrial quadrupedal knuckle-walking species, more similar to extant chimpanzees.<\/span><\/li>\n<\/ol>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Most research supports the first theory of an arboreal LCA based on skeletal morphology of early hominin genera that demonstrate adaptations for climbing but not for knuckle-walking. This would mean that both humans and chimpanzees can be considered \u201cderived\u201d in terms of locomotion since chimpanzees would have independently evolved knuckle-walking.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are many current ideas regarding selective pressures that would lead to early hominins adapting upright posture and locomotion. Many of these selective pressures, as we have seen in the previous section, coincide with a shift in environmental conditions, supported by paleoenvironmental data. In general, however, it appears that, like extant great apes, early hominins thrived in forested regions with dense tree coverage, which would indicate an arboreal lifestyle. As the environmental conditions changed and a savannah\/grassland environment became more widespread, the tree cover would become less dense, scattered, and sparse such that bipedalism would become more important.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are several proposed selective pressures for bipedalism:<\/span><\/p>\n<ol>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\"><strong>E<\/strong><strong>nergy conservation:<\/strong> Modern bipedal humans conserve more energy than extant chimpanzees, which are predominantly knuckle-walking quadrupeds when walking over land. While chimpanzees, for instance, are faster than humans terrestrially, they expend large amounts of energy being so. Adaptations to bipedalism include \u201cstacking\u201d the majority of the weight of the body over a small area around the center of gravity (i.e., the head is above the chest, which is above the pelvis, which is over the knees, which are above the feet). This reduces the amount of muscle needed to be engaged during locomotion to \u201cpull us up\u201d and allows us to travel longer distances expending far less energy.<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\"><strong>T<\/strong><strong>hermoregulation:<\/strong> Less surface area (i.e., only the head and shoulders) is exposed to direct sunlight during the hottest parts of the day (i.e., midday). This means that the body has less need to employ additional \u201ccooling\u201d mechanisms such as sweating, which additionally means less water loss.<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Bipedalism <\/strong><span style=\"text-decoration: underline\">(Freeing of Hands)<\/span><strong>: <\/strong>This method of locomotion freed up our ancestors\u2019 hands such that they could more easily gather food and carry tools or infants. This further enabled the use of hands for more specialized adaptations associated with the manufacturing and use of tools.<\/span><\/li>\n<\/ol>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">These selective pressures are not mutually exclusive. Bipedality could have evolved from a combination of these selective pressures, in ways that increased the chances of early hominin survival.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Skeletal Adaptations for Bipedalism<\/strong><\/span><\/h3>\n<figure style=\"width: 405px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image10-1.png\" alt=\"A full human skeleton and gorilla skeleton standing in upright positions next to each other.\" width=\"405\" height=\"452\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.6: Compared to gorillas (right) and other apes, humans (left) have highly specialized adaptations to facilitate bipedal locomotion. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Primatenskelett-drawing.jpg\">Skeleton of human (1) and gorilla (2), unnaturally sketched<\/a> by unknown from Brehms Tierleben, Small Edition 1927 is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Humans have highly specialized adaptations to facilitate obligate bipedalism (Figure 9.6). Many of these adaptations occur within the soft tissue of the body (e.g., muscles and tendons). However, when analyzing the paleoanthropological record for evidence of the emergence of bipedalism, all that remains is the fossilized bone. Interpretations of locomotion are therefore often based on comparative analyses between fossil remains and the skeletons of extant primates with known locomotor behaviors. These adaptations occur throughout the skeleton and are summarized in Figure 9.7.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The majority of these adaptations occur in the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1336\"><strong>postcranium<\/strong><\/a> <span style=\"background-color: #ff99cc\">(the skeleton from below the head)<\/span> and are outlined in Figure 9.7. In general, these adaptations allow for greater stability and strength in the lower limb, by allowing for more shock absorption, for a larger surface area for muscle attachment, and for the \u201cstacking\u201d of the skeleton directly over the center of gravity to reduce energy needed to be kept upright. These adaptations often mean less flexibility in areas such as the knee and foot.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">However, these adaptations come at a cost. Evolving from a nonobligate bipedal ancestor means that the adaptations we have are evolutionary compromises. For instance, the valgus knee (angle at the knee) is an essential adaptation to balance the body weight above the ankle during bipedal locomotion. However, the strain and shock absorption at an angled knee eventually takes its toll. For example, runners often experience joint pain. Similarly, the long neck of the femur absorbs stress and accommodates for a larger pelvis, but it is a weak point, resulting in hip replacements being commonplace among the elderly, especially in cases where the bone additionally weakens through osteoporosis. Finally, the S-shaped curve in our spine allows us to stand upright, relative to the more curved C-shaped spine of an LCA. Yet the weaknesses in the curves can lead to pinching of nerves and back pain. Since many of these problems primarily are only seen in old age, they can potentially be seen as an evolutionary compromise.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Despite relatively few postcranial fragments, the fossil record in early hominins indicates a complex pattern of emergence of bipedalism. Key features, such as a more anteriorly placed foramen magnum, are argued to be seen even in the earliest discovered hominins, indicating an upright posture (Dart 1925). Some early species appear to have a mix of ancestral (arboreal) and derived (bipedal) traits, which indicates a mixed locomotion and a more <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1402\">mosaic evolution<\/a><\/strong> of the trait. Some early hominins appear to, for instance, have bowl-shaped pelvises (hip bones) and angled femurs suitable for bipedalism but also have retained an opposable <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1338\">hallux<\/a><\/strong> (big toe) or curved fingers and longer arms (for arboreal locomotion). These mixed morphologies may indicate that earlier hominins were not fully obligate bipeds and thus thrived in mosaic environments.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Yet the associations between postcranial and the more diagnostic cranial fossils and bones are not always clear, muddying our understanding of the specific species to which fossils belong (Grine et al. 2022).<\/span><\/p>\n<div align=\"left\">\n<table class=\"grid\">\n<caption>\n<p class=\"import-Normal\" style=\"text-align: left\"><span style=\"color: #000000\">Figure 9.7: Skeletal comparisons between modern humans (obligate bipeds) and nonobligate bipeds (e.g., chimpanzees). Credit: <a class=\"rId34\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\">Skeletal comparisons between modern humans and <\/a><a class=\"rId35\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\">nonobligate<\/a><a class=\"rId36\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\"> bipeds (Figure 9.6)<\/a> original to <a class=\"rId37\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a class=\"rId38\" style=\"color: #000000\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/span><\/p>\n<\/caption>\n<thead>\n<tr>\n<td style=\"width: 97.998px\"><strong>Region<\/strong><\/td>\n<td style=\"width: 106.992px\"><strong>Feature<\/strong><\/td>\n<td style=\"width: 366.992px\"><strong>Obligate Biped (H. sapiens)<\/strong><\/td>\n<td style=\"width: 310px\"><strong>Nonobligate Biped<\/strong><\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr>\n<td style=\"width: 97.998px\">Cranium<\/td>\n<td style=\"width: 106.992px\">Position of the foramen magnum<\/td>\n<td style=\"width: 366.992px\">Positioned inferiorly (immediately under the cranium) so that the head rests on top of the vertebral column for balance and support (head is perpendicular to the ground).<\/td>\n<td style=\"width: 310px\">Posteriorly positioned (to the back of the cranium). Head is positioned parallel to the ground.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Body proportions<\/td>\n<td style=\"width: 366.992px\">Shorter upper limb (not used for locomotion).<\/td>\n<td style=\"width: 310px\">Longer upper limbs (used for locomotion).<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Spinal curvature<\/td>\n<td style=\"width: 366.992px\">S-curve due to pressure exerted on the spine from bipedalism (lumbar lordosis).<\/td>\n<td style=\"width: 310px\">C-curve.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Vertebrae<\/td>\n<td style=\"width: 366.992px\">Robust lumbar (lower-back) vertebrae (for shock absorbance and weight bearing). Lower back is more flexible than that of apes as the hips and trunk swivel when walking (weight transmission).<\/td>\n<td style=\"width: 310px\">Gracile lumbar vertebrae compared to those of modern humans.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Pelvis<\/td>\n<td style=\"width: 366.992px\">Shorter, broader, bowl-shaped pelvis (for support); very robust. Broad sacrum with large sacroiliac joint surfaces.<\/td>\n<td style=\"width: 310px\">Longer, flatter, elongated ilia; more narrow and gracile; narrower sacrum; relatively smaller sacroiliac joint surface.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Lower limb<\/td>\n<td style=\"width: 366.992px\">In general, longer, more robust lower limbs and more stable, larger joints.<\/p>\n<ul>\n<li style=\"font-weight: 400\">Large femoral head and longer neck (absorbs more stress and increases the mechanical advantage).<\/li>\n<li style=\"font-weight: 400\">Valgus knee, in which the angle of the knee positions it over the ankle and keeps the center of gravity balanced over the stance leg during stride cycle (shock absorbance).<\/li>\n<li style=\"font-weight: 400\">Distal tibia (lower leg) of humans has a large medial malleolus for stability.<\/li>\n<\/ul>\n<\/td>\n<td style=\"width: 310px\">In general, smaller, more gracile limbs with more flexible joints.<\/p>\n<ul>\n<li style=\"font-weight: 400\">Femoral neck is smaller in comparison to modern humans and shorter.<\/li>\n<li style=\"font-weight: 400\">The legs bow outward, and there is no valgus angle of the knee (no \u201cknock knees\u201d).<\/li>\n<li style=\"font-weight: 400\">The distal tibia in chimpanzees is trapezoid (wider anteriorly) for climbing and allows more flexibility.<\/li>\n<\/ul>\n<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Foot<\/td>\n<td style=\"width: 366.992px\">Rigid, robust foot, without a midtarsal break.<\/p>\n<p>Nonopposable and large, robust big toe (for push off while walking) and large heel for shock absorbance.<\/td>\n<td style=\"width: 310px\">Flexible foot, midtarsal break present (which allows primates to lift their heels independently from their feet), opposable big toe for grasping.<\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<p class=\"import-Normal\"><span style=\"color: #000000;background-color: #ccffcc\">It is also worth noting that, while not directly related to bipedalism per se, other postcranial adaptations are evident in the hominin fossil record from some of the earlier hominins. For instance, the hand and finger morphologies of many of the earliest hominins indicate adaptations consistent with arboreality. These include longer hands, more curved metacarpals and phalanges (long bones in the hand and fingers, respectively), and a shorter, relatively weaker thumb. This allows for gripping onto curved surfaces during locomotion. The earliest hominins appear to have mixed morphologies for both bipedalism and arborealism. However, among Australopiths (members of the genus, Australopithecus), there are indications for greater reliance on bipedalism as the primary form of locomotion. Similarly, adaptations consistent with tool manufacture (shorter fingers and a longer, more robust thumb, in contrast to the features associated with arborealism) have been argued to appear before the genus <em>Homo<\/em>.<\/span><\/p>\n<div class=\"textbox shaded\">\n<h2>Special Topic: Fear of Snakes \u2014 A Cultural or Biological Response\/Adaptation?<\/h2>\n<figure style=\"width: 245px\" class=\"wp-caption alignright\"><img src=\"https:\/\/www.researchgate.net\/profile\/David-Penning\/publication\/315305069\/figure\/fig2\/AS:613947958902785@1523388008174\/Constriction-coil-postures-of-kingsnakes-and-ratsnakes-Typical-constriction-coil.png\" alt=\"Constriction coil postures of kingsnakes and ratsnakes. Typical constriction coil postures in a kingsnake, Lampropeltis getula (92 g; A), and a ratsnake, Pantherophis guttatus (86 g; B). Both snakes were constricting similarly sized mice, Mus musculus (12 g). The relative prey mass was 13% for the kingsnake and 13.9% for the ratsnake.\u00a0\" width=\"245\" height=\"245\" \/><figcaption class=\"wp-caption-text\">Constriction coil postures of kingsnakes and ratsnakes. source: https:\/\/www.researchgate.net\/figure\/Constriction-coil-postures-of-kingsnakes-and-ratsnakes-Typical-constriction-coil_fig2_315305069<\/figcaption><\/figure>\n<p>It is suggested that primates have three major predators: raptors, felines, and snakes; however, many studies show that of these carnivores, snakes were one of the first that mammals had to contend with alongside dinosaurs, as felines and raptors evolved at a much slower pace than their reptilian competition. Herpetologists trace the evolution of constricting snakes to about 100 million years ago, and by the time mammals arrived around 75 million years ago, constrictors were\u00a0 already well established as a formidable threat (Greene, 2017). \u00a0Both co-existed for millennia and each sustained selective pressures requiring them to evolve specific traits to survive. When venomous snakes eventually emerged 55 to 65 million years ago, they posed yet an additional threat to proto-primates as they required less distance for the predator to kill (2017). Alongside camouflage and silent movement techniques, it was the development of the snake\u2019s hollow fangs through which to deliver venom that was most transformative to primate evolution. As such, primates evolved their pre-conscious attention, and visual acuity to cope with this new threat; therefore, while snakes were adapting morphologically to feed themselves, they were unwittingly teaching proto-primates valuable lessons in predator detection and reacting appropriately in order to survive.<\/p>\n<p>In a 2009 Harvard University study, Lynne A. Isbell hypothesizes that envenoming snakes are linked to being directly responsible for the origins of the evolving complex brains and superior visual capacity in the lineage of anthropoids leading to humans (Isbell, 2009). Forward-facing eyes for binocular vision, depth perception, enhanced visual acuity, stereoscopic and trichromatic colour vision, all traits necessary for snake detection; and the quick motor responses from the primate\u2019s fight, flight, or freeze defence mechanism to circumvent a snake\u2019s squeeze or bite. Numerous laboratory studies show that humans and primates both sense and visually detect snakes more rapidly than other threatening stimuli (Van Le Et al., 2013). These experiments show that snakes elicited the strongest, fastest responses (Van Le Et al., 2013). This is known as \u2018Snake Detection Theory\u2019 and is the evolution of the primate\u2019s complex brain, visual acuity, and rapid motor responses towards snakes in its environment that are the adaptations needed to live successfully as arboreal beings. It is not fortuitous then, that primates that never coexisted with venomous snakes, such as lemurs in Madagascar, have less visual acuity, better olfaction and smaller brains. Within Isbell\u2019s work, a collaborative study by a group of neuroscientists tested this hypothesis and found that, indeed, there is higher neural firing and activity in multiple areas of the primate brain, notably in the pulvinar, a region\u00a0 responsible for visual attention and oculomotor behaviour (Isbell, L., 2009).<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 509px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/www.worldhistory.org\/img\/r\/p\/750x750\/4941.jpg?v=1747098791\" alt=\"Apophis Defeated\" width=\"509\" height=\"309\" \/><figcaption class=\"wp-caption-text\">Apophis being killed by the knife-wielding cat of Heliopolis, an agent of Re;245 Tomb of Inherkhau, Deir el-Medina (TT 359). Note the very large eye of the snake, the source of its malevolent stare or \u201cevil eye.\u201d source: Weeks, Kent R. (2005) The Illustrated Guide to Luxor: Tombs, Temples and Museums, The American University in Cairo Press, Cairo.<\/figcaption><\/figure>\n<p>Today, the fear of snakes is widespread in humans, often shown through avoidance and disgust. A study in <em data-start=\"197\" data-end=\"244\">The Journal of Ethnobiology and Ethnomedicine<\/em> notes that snakes are over-hunted and excluded from conservation efforts worldwide (Cer\u00edaco, 2012). While cultural factors shape our sentiments, instinct also plays a role\u2014such as the developed avoidance behaviors toward threats like snakes. This blend of instinct and cultural influence is not only seen in behavior but also deeply embedded in the stories we tell. Many cultures depict mythological snakes as harbingers of death or chaos. In the Bible, Satan becomes a snake to tempt Eve. Norse mythology features J\u00f6rmungandr, the world serpent who signals the apocalypse. Egyptian myth tells of Apophis, who battles the sun god Ra nightly. Though sources vary, these myths consistently portray snakes as threats. As such, the widespread fear of snakes may reflect both evolutionary and cultural influences. Understood as an adaptive response inherited from primate ancestors\u2014who developed avoidance behaviors toward potentially dangerous stimuli\u2014and reinforced through myths and religious narratives, the enduring presence of snakes as potent figures of fear across human societies and primate groups highlights the complex intertwining of instinct and cultural meaning in shaping human behavior.<\/p>\n<\/div>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Early Hominins: Sahelanthropus and Orrorin<\/strong><\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">We see evidence for bipedalism in some of the earliest fossil hominins, dated from within our estimates of our divergence from chimpanzees. These hominins, however, also indicate evidence for arboreal locomotion.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The earliest dated hominin find (between 6 mya and 7 mya, based on radiometric dating of volcanic tufts) has been argued to come from Chad and is named <strong><em>Sahelanthropus tchadensis<\/em> <\/strong>(Figure 9.8; Brunet et al. 1995). The initial discovery was made in 2001 by Ahounta Djimdoumalbaye and announced in <em>Nature<\/em> in 2002 by a team led by French paleontologist Michel Brunet. The find has a small cranial capacity (360 cc) and smaller canines than those in extant great apes, though they are larger and pointier than those in humans. <span style=\"background-color: #ffff00\">This implies strongly that, over evolutionary time, the need for display and dominance among males has reduced, as has our sexual dimorphism<\/span>. A short cranial base and a foramen magnum that is more humanlike in positioning have been argued to indicate upright walking.<\/span><\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 640px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-289\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.82.jpg\" alt=\"Four views of a beige-colored skull are seen against a black background.\" width=\"640\" height=\"640\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.8: Sahelanthropus tchadensis exhibits a set of derived features, including a long, low cranium; a small, ape-sized braincase; and relatively reduced prognathism. Credit: aa <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 anterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 posterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 inferior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; and d <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 lateral left view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Initially, the inclusion of <em>Sahelanthropus<\/em> in the hominin family was debated by researchers, since the evidence for bipedalism is based on cranial evidence alone, which is not as convincing as postcranial evidence. Yet, a femur (thigh bone) and ulnae (upper arm bones) thought to belong to <em>Sahelanthropus<\/em> was discovered in 2001 (although not published until 2022). These bones may support the idea that the hominin was in fact a terrestrial biped with arboreal capabilities and behaviors (Daver et al. 2022).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Orrorin tugenensis<\/em><\/strong> (Orrorin meaning \u201coriginal man\u201d), dated to between 6 mya and 5.7 mya, was discovered near Tugen Hills in Kenya in 2000. Smaller <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1340\">cheek teeth<\/a><\/strong> (molars and premolars) than those in even more recent hominins, thick enamel, and reduced, but apelike, canines characterize this species. This is the first species that clearly indicates adaptations for bipedal locomotion, with fragmentary leg, arm, and finger bones having been found but few cranial remains. One of the most important elements discovered was a proximal femur, BAR 1002'00. The femur is the thigh bone, and the proximal part is that which articulates with the pelvis; this is very important for studying posture and locomotion. This femur indicates that <em>Ororrin<\/em> was bipedal, and recent studies suggest that it walked in a similar way to later <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1404\">Pliocene<\/a><\/strong> hominins. Some have argued that features of the finger bones suggest potential tool-making capabilities, although many researchers argue that these features are also consistent with climbing.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Early Hominins: The Genus <em>Ardipithecus<\/em><\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Another genus, <em>Ardipithecus<\/em>, is argued to be represented by at least two species: <em>Ardipithecus (Ar.) ramidus <\/em>and <em>Ar. kadabba<\/em>.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Ardipithecus ramidus<\/em><\/strong> (\u201cramid\u201d means root in the Afar language) is currently the best-known of the earliest hominins (Figure 9.9). Unlike <em>Sahelanthropus<\/em> and<em> Orrorin<\/em>, this species has a large sample size of over 110 specimens from Aramis alone. Dated to 4.4 mya, <em>Ar. ramidus<\/em> was found in Ethiopia (in the Middle Awash region and in Gona). This species was announced in 1994 by American palaeoanthropologist Tim White, based on a partial female skeleton nicknamed \u201cArdi\u201d (ARA-VP-6\/500; White et al. 1994). Ardi demonstrates a mosaic of ancestral and derived characteristics in the postcrania. For instance, she had an opposable big toe (hallux), similar to chimpanzees (i.e., more ancestral), which could have aided in climbing trees effectively. However, the pelvis and hip show that she could walk upright (i.e., it is derived), supporting her hominin status. A small brain (300 cc to 350 cc), midfacial projection, and slight prognathism show retained ancestral cranial features, but the cheek bones are less flared and robust than in later hominins.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 706px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-1310\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.9-scaled-1.jpg\" alt=\"Skull cast and partial skeleton with photographs of some bones and line drawings of others.\" width=\"706\" height=\"453\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.9a and b: Researchers believe that Ardipithecus ramidus was able to walk upright, although not as efficiently as later humans. It possessed the musculature required for tree climbing, and while moving quadrupedally, it likely placed weight on the palms of the hands rather than on the knuckles. Credit: a. <a class=\"rId61\" href=\"https:\/\/boneclones.com\/product\/ardipithecus-ramidus-skull-BH-039\">Ardipithecus ramidus Skull<\/a> by <a class=\"rId62\" href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a> is used by permission and available here under a <a class=\"rId63\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>; b. <a class=\"rId64\" href=\"https:\/\/boneclones.com\/product\/ardipithecus-ramidus-skull-BH-039\">Artist\u2019s rendition of \u201cArdi\u201d skeleton<\/a> by <a class=\"rId65\" href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a> is used by permission and available here under a <a class=\"rId66\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong><em>Ardipithecus kadabba<\/em><\/strong> (the species name means \u201coldest ancestor\u201d in the Afar language) is known from localities on the western margin of the Middle Awash region, the same locality where <em>Ar. ramidus<\/em> has been found. Specimens include mandibular fragments and isolated teeth as well as a few postcranial elements from the Asa Koma (5.5 mya to 5.77 mya) and Kuseralee Members (5.2 mya), well-dated and understood (but temporally separate) volcanic layers in East Africa. This species was discovered in 1997 by paleoanthropologist Dr. Yohannes Haile-Selassie. Originally these specimens were referred to as a subspecies of <em>Ar. ramidus<\/em>. In 2002, six teeth were discovered at Asa Koma and the dental-wear patterns confirmed that this was a distinct species, named <em>Ar. kadabba,<\/em> in 2004. One of the postcranial remains recovered included a 5.2 million-year-old toe bone that demonstrated features that are associated with toeing off (pushing off the ground with the big toe leaving last) during walking, a characteristic unique to bipedal walkers. However, the toe bone was found in the Kuseralee Member, and therefore some doubt has been cast by researchers about its association with the teeth from the Asa Koma Member.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"background-color: #ff99cc\"><strong><span style=\"color: #000000\">Bipedal Trends in Early Hominins: Summary<\/span><\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff99cc\">Trends toward bipedalism are seen in our earliest hominin finds. However, many specimens also indicate retained capabilities for climbing. Trends include a larger, more robust hallux; a more compact foot, with an arch; a robust, long femur, angled at the knee; a robust tibia; a bowl-shaped pelvis; and a more anterior foramen magnum. While the level of bipedality in <em>Salehanthropus<\/em> <em>tchadenisis<\/em> is debated since there are few fossils and no postcranial evidence, <em>Orrorin tugenensis<\/em> and <em>Ardipithecus<\/em> <em>k<\/em><em>adabba <\/em>show clear indications of some of these bipedal trends. However, some retained ancestral traits, such as an opposable hallux in <em>Ardipithecus<\/em>, indicate some retention in climbing ability.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Derived Adaptations: Early Hominin Dention<\/span><\/h2>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">The Importance of Teeth<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Teeth are abundant in the fossil record, primarily because they are already highly mineralized as they are forming, far more so than even bone. Because of this, teeth preserve readily. And, because they preserve readily, they are well-studied and better understood than many skeletal elements. In the sparse hominin (and primate) fossil record, teeth are, in some cases, all we have.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Teeth also reveal a lot about the individual from whom they came. We can tell what they evolved to eat, to which other species they may be closely related, and even, to some extent, the level of sexual dimorphism, or general variability, within a given species. This is powerful information that can be contained in a single tooth. With a little more observation, the wearing patterns on a tooth can tell us about the diet of the individual in the weeks leading up to its death. Furthermore, the way in which a tooth is formed, and the timing of formation, can reveal information about changes in diet (or even mobility) over infancy and childhood, using isotopic analyses. When it comes to our earliest hominin relatives, this information is vital for understanding how they lived.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The purpose of comparing different hominin species is to better understand the functional morphology as it applies to dentition. In this, we mean that the morphology of the teeth or masticatory system (which includes jaws) can reveal something about the way in which they were used and, therefore, the kinds of foods these hominins ate. When comparing the features of hominin groups, it is worth considering modern analogues (i.e., animals with which to compare) to make more appropriate assumptions about diet. In this way, hominin dentition is often compared with that of chimpanzees and gorillas (our close ape relatives), as well as with that of modern humans.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The most divergent group, however, is humans. Humans around the world have incredibly varied diets. Among hunter-gatherers, it can vary from a honey- and plant-rich diet, as seen in the Hadza in Tanzania, to a diet almost entirely reliant on animal fat and protein, as seen in Inuits in polar regions of the world. We are therefore considered generalists, more general than the largely <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1342\"><strong>frugivorous<\/strong><\/a> (fruit-eating) chimpanzee or the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1344\"><strong>folivorous<\/strong><\/a> (foliage-eating) gorilla, as discussed in Chapter 5.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">One way in which all humans are similar is our reliance on the processing of our food. We cut up and tear meat with tools using our hands, instead of using our front teeth (incisors and canines). We smash and grind up hard seeds, instead of crushing them with our hind teeth (molars). This means that, unlike our ape relatives, we can rely more on developing tools to navigate our complex and varied diets. <span style=\"text-decoration: underline\">(We could say)<\/span> Our brain, therefore, is our primary masticatory organ. Evolutionarily, our teeth have reduced in size and our faces are flatter, or more <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1346\">orthognathic<\/a>, <\/strong>partially in response to our increased reliance on our hands and brain to process food. Similarly, a reduction in teeth and a more generalist dental morphology could also indicate an increase in softer and more variable foods, such as the inclusion of more meat. <span style=\"background-color: #ffff00\">These trends begin early on in our evolution.<\/span> The link has been made between some of the earliest evidence for stone tool manufacture, the earliest members of our genus, and the features that we associate with these specimens.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">General Dental Trends in Early Hominins<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Several trends are visible in the dentition of early hominins. However, all tend to have the same <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1348\"><strong>dental formula<\/strong><\/a>. The dental formula tells us how many of each tooth type are present in each quadrant of the mouth. Going from the front of the mouth, this includes the square, flat <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1160\">incisors<\/a><\/strong>; the pointy <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1166\">canines<\/a><\/strong>; the small, flatter <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1350\"><strong>premolars<\/strong><\/a>; and the larger hind <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1164\">molars<\/a><\/strong>. In many primates, from Old World monkeys to great apes, the typical dental formula is 2:1:2:3. This means that if we divide the mouth into quadrants, each has two incisors, one canine, two premolars, and three molars. The eight teeth per quadrant total 32 teeth in all (although some humans have fewer teeth due to the absence of their wisdom teeth, or third molars).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 380px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image24.png\" alt=\"Anterior view of the lower face of a person showing their teeth.\" width=\"380\" height=\"253\" \/><figcaption class=\"wp-caption-text\">Figure 9.10: In humans, our canines are often a similar size to our incisors. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Adult_human_teeth.jpg\">Adult human teeth<\/a> by <a href=\"https:\/\/www.genusfotografen.se\/\">Genusfotografen<\/a> (Tomas Gunnarsson) through <a href=\"https:\/\/wikimedia.se\/\">Wikimedia Sverige<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p><span style=\"color: #000000\">The morphology of the individual teeth is where we see the most change. Among primates, large incisors are associated with food procurement or preparation (such as biting small fruits), while small incisors indicate a diet that may contain small seeds or leaves (where the preparation is primarily in the back of the mouth). Most hominins have relatively large, flat, vertically aligned incisors that <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1396\">occlude<\/a> <\/strong>(touch) relatively well, forming a \u201cbite.\u201d This differs from, for instance, the orangutan, whose teeth stick out (i.e.<em>,<\/em> are <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1352\"><strong>procumbent<\/strong><\/a>).<\/span><\/p>\n<p>&nbsp;<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">While the teeth are often aligned with diet, the canines may be misleading in that regard. We tend to associate pointy, large canines with the ripping required for meat, and the reduction (or, in some animals, the absence) of canines as indicative of herbivorous diets. In humans, our canines are often a similar size to our incisors and therefore considered <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1398\">incisiform<\/a><\/strong> (Figure 9.10). However, our closest relatives all have very long, pointy canines, particularly on their upper dentition. This is true even for the gorilla, which lives almost exclusively on plants. The canines in these instances reveal more about social structure and sexual dimorphism than diet, as large canines often signal <span style=\"background-color: #ffff00\">dominance<\/span>.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Early on in human evolution, we see a reduction in canine size. <em>Sahelanthropus tchadensis<\/em> and <em>Orrorin tugenensis<\/em> both have smaller canines than those in extant great apes, yet the canines are still larger and pointier than those in humans or more recent hominins. <span style=\"background-color: #ffff00\">This implies strongly that, over evolutionary time, the need for display and dominance among males has reduced, as has our sexual dimorphism.<\/span> In <em>Ardipithecus ramidus<\/em>, there is no obvious difference between male and female canine size, yet they are still slightly larger and pointier than in modern humans. This implies a less sexually dimorphic social structure in the earlier hominins relative to modern-day chimpanzees and gorillas.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Along with a reduction in canine size is the reduction or elimination of a canine <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1172\">diastema<\/a>:<\/strong> a gap between the teeth on the mandible that allows room for elongated teeth on the maxilla to \u201cfit\u201d in the mouth. Absence of a diastema is an excellent indication of a reduction in canine size. In animals with large canines (such as baboons), there is also often a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1354\"><strong>honing P3<\/strong><\/a>, where the first premolar (also known as P3 for evolutionary reasons) is triangular in shape, \u201csharpened\u201d by the extended canine from the upper dentition. This is also seen in some early hominins: <em>Ardipithecus<\/em>, for example, has small canines that are almost the same height as its incisors, although still larger than those in recent hominins.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The hind dentition, such as the bicuspid (two cusped) premolars or the much larger molars, are also highly indicative of a generalist diet in hominins. Among the earliest hominins, the molars are larger than we see in our genus, increasing in size to the back of the mouth and angled in such a way from the much smaller anterior dentition as to give these hominins a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1356\"><strong>parabolic<\/strong><\/a> (V-shaped) dental arch. This differs from our living relatives and some early hominins, such as <em>Sahelanthropus<\/em>, whose molars and premolars are relatively parallel between the left and right sides of the mouth, creating a U-shape.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Among more recent early hominins, the molars are larger than those in the earliest hominins and far larger than those in our own genus, <em>Homo.<\/em> Large, short molars with thick <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1358\"><strong>enamel<\/strong><\/a> allowed our early cousins to grind fibrous, coarse foods, such as sedges, which require plenty of chewing. This is further evidenced in the low <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1176\">cusps<\/a>,<\/strong> or ridges, on the teeth, which are ideal for chewing. In our genus, the hind dentition is far smaller than in these early hominins. Our teeth also have medium-size cusps, which allow for both efficient grinding and tearing\/shearing meats.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Understanding the dental morphology has allowed researchers to extrapolate very specific behaviors of early hominins. It is worth noting that while teeth preserve well and are abundant, a slew of other morphological traits additionally provide evidence for many of these hypotheses. Yet there are some traits that are ambiguous. For instance, while there are definitely high levels of sexual dimorphism in <em>Au. afarensis<\/em>, discussed in the next section, the canine teeth are reduced in size, implying that while canines may be useful indicators for sexual dimorphism, it is also worth considering other evidence.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff99cc\">In summary, trends among early hominins include a reduction in procumbency, reduced hind dentition (molars and premolars), a reduction in canine size (more incisiform with a lack of canine diastema and honing P3), flatter molar cusps, and thicker dental enamel. All early hominins have the ancestral dental formula of 2:1:2:3. These trends are all consistent with a generalist diet, incorporating more fibrous foods.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Special Topic: Contested Species<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Many named species are highly debated and argued to have specimens associated with a more variable <em>Au. afarensis <\/em>or <em>Au. anamensis<\/em> species. Sometimes these specimens are dated to times when, or found in places in which, there are \u201cgaps\u201d in the palaeoanthropological record. These are argued to represent chronospecies or variants of <em>Au. afarensis<\/em>. However, it is possible that, with more discoveries, the distinct species types will hold.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><strong><em>Australopithecus bahrelghazali<\/em><\/strong> is dated to within the time period of <em>Au. afarensi<\/em>s (3.6 mya; Brunet et al. 1995) and was the first Australopithecine to be discovered in Chad in central Africa. Researchers argue that the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1360\">holotype<\/a><\/strong>, whom discoverers have named \u201cAbel,\u201d falls under the range of variation of <em>Au. afarensis<\/em> and therefore that <em>A. bahrelghazali<\/em> does not fall into a new species (Lebatard et al. 2008). If \u201cAbel\u201d is a member of <em>Au. afarensis<\/em>, the geographic range of the species would be greatly extended.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">On a different note, <strong><em>Australopithecus <\/em><\/strong><strong><em>deyiremada<\/em><\/strong> (meaning \u201cclose relative\u201d in the Ethiopian language of Afar) is dated to 3.5 mya to 3.3 mya and is based on fossil mandible bones discovered in 2011 in Woranso-Mille (in the Afar region of Ethiopia) by Yohannes Haile-Selassie, an Ethiopian paleoanthropologist (Haile-Selassie et al. 2019). The discovery indicated, in contrast to <em>Au. afarensis<\/em>, smaller teeth with thicker enamel (potentially suggesting a harder diet) as well as a larger mandible and more projecting cheekbones. This find may be evidence that more than one closely related hominin species occupied the same region at the same temporal period (Haile-Selassie et al. 2015; Spoor 2015) or that other <em>Au. afarensis<\/em> specimens have been incorrectly designated. However, others have argued that this species has been prematurely identified and that more evidence is needed before splitting the taxa, since the variation appears subtle and may be due to slightly different niche occupations between populations over time.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><strong><em>Australopithecus garhi<\/em><\/strong> is another species found in the Middle Awash region of Ethiopia. It is currently dated to 2.5 mya (younger than <em>Au. afarensis<\/em>). Researchers have suggested it fills in a much-needed temporal \u201cgap\u201d between hominin finds in the region, with some anatomical differences, such as a relatively large cranial capacity (450 cc) and larger hind dentition than seen in other gracile Australopithecines. Similarly, the species has been argued to have longer hind limbs than <em>Au. afarensis<\/em>, although it was still able to move arboreally (Asfaw et al. 1999). However, this species is not well documented or understood and is based on only several fossil specimens. More astonishingly, crude stone tools resembling Oldowan (which will be described later) have been found in association with <em>Au. garhi<\/em>. While lacking some of the features of the Oldowan, this is one of the earliest technologies found in direct association with a hominin.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><strong><em>Kenyanthopus<\/em><\/strong><strong><em> platyops<\/em><\/strong> (the name \u201cplatyops\u201d refers to its flatter-faced appearance) is a highly contested genus\/species designation of a specimen (KNM-WT 40000) from Lake Turkana in Kenya, discovered by Maeve Leakey in 1999 (Figure 9.11). Dated to between 3.5 mya and 3.2 mya, some have suggested this specimen is an <em>Australopithecus<\/em>, perhaps even <em>Au.<\/em> <em>afarensis<\/em> (with a brain size which is difficult to determine, yet appears small), while still others have placed this specimen in <em>Homo <\/em>(small dentition and flat-orthognathic face). While taxonomic placing of this species is quite divided, the discoverers have argued that this species is ancestral to <em>Homo<\/em>, in particular to <em>Homo <\/em><em>ruldolfensis<\/em> (Leakey et al. 2001). Some researchers have additionally associated the earliest tool finds from Lomekwi, Kenya, temporally (3.3 mya) and in close geographic proximity to this specimen.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 579px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-292 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.11.jpg\" alt=\"Four views of an ancient skull are shown on a black background.\" width=\"579\" height=\"579\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.11: This specimen, KNM WT 40000 (Kenyanthopus platyops), has small detention, a small brain case, and a relatively flat face. Its genus\/species designation remains contested. Credit: a. <a class=\"rId76\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId77\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 anterior view<\/a> by \u00a9<a class=\"rId78\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId79\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId80\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId81\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId82\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 superior view<\/a> by \u00a9<a class=\"rId83\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId84\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId85\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a class=\"rId86\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId87\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 lateral left view<\/a> by \u00a9<a class=\"rId88\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId89\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId90\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; d. <a class=\"rId91\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId92\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 inferior view<\/a> by \u00a9<a class=\"rId93\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId94\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId95\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">The Genus <em>Australopithecus<\/em><br \/>\n<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The Australopithecines are a diverse group of hominins, comprising various species. <em>Australopithecus<\/em> is the given group or genus name. It stems from the Latin word <em>Australo<\/em>, meaning \u201csouthern,\u201d and the Greek word <em>pithecus,<\/em> meaning \u201cape.\u201d Within this section, we will outline these differing species\u2019 geological and temporal distributions across Africa, unique derived and\/or shared traits, and importance in the fossil record.<\/span><\/p>\n<figure style=\"width: 381px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image37-2.png\" alt=\"A skull has a pronounced sagittal crest, flaring cheekbones, and large hind teeth.\" width=\"381\" height=\"585\" \/><figcaption class=\"wp-caption-text\">Figure 9.12: Robust Australopithecines such as Paranthropus boisei had large molars and chewing muscles. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Paranthropus_boisei_skull.jpg\">Paranthropus boisei skull<\/a> by Durova is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/deed.en\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Between 3 mya and 1 mya, there seems to be differences in dietary strategy between different species of hominins designated as Australopithecines. A pattern of larger posterior dentition (even relative to the incisors and canines in the front of the mouth), thick enamel, and cranial evidence for extremely large chewing muscles is far more pronounced in a group known as the robust australopithecines. This pattern is extreme<span style=\"text-decoration: underline\">ly<\/span> relative to their earlier contemporaries or predecessors, the gracile australopithecines<strong>,<\/strong> and is certainly larger than those seen in early <em>Homo<\/em>, which emerged during this time. This pattern of incredibly large hind dentition (and very small anterior dentition) has led people to refer to robust australopithecines as <strong>megadont<\/strong> hominins (Figure 9.12).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Because of these differences, this section has been divided into \u201cgracile\u201d and \u201crobust\u201d Australopithecines, highlighting the morphological differences between the two groups (which many researchers have designated as separate genera: <em>Australopithecus<\/em> and <em>Paranthropus<\/em>, respectively) and then focusing on the individual species. It is worth noting, however, that not all researchers accept these clades as biologically or genetically distinct, with some researchers insisting that the relative gracile and robust features found in these species are due to parallel evolutionary events toward similar dietary niches.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Despite this genus\u2019 ancestral traits and small cranial capacity, all members show evidence of bipedal locomotion. It is generally accepted that <em>Australopithecus <\/em>species display varying degrees of arborealism along with bipedality.<\/span><\/p>\n<h3 class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Gracile Australopithecines<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">This section describes individual species from across Africa. These species are called \u201c<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1406\">gracile<\/a> <\/strong>australopithecines\u201d because of their smaller and less robust features compared to the divergent \u201c<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1408\"><strong>robust<\/strong><\/a>\u201d group. Numerous Australopithecine species have been named, but some are only based on a handful of fossil finds, whose designations are controversial.<\/span><\/p>\n<h4 class=\"import-Normal\"><em><span style=\"color: #000000\">East African Australopithecines<\/span><\/em><\/h4>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">East African Australopithecines are found throughout the EARS, and they include the earliest species associated with this genus. Numerous fossil-yielding sites, such as Olduvai, Turkana, and Laetoli, have excellent, datable stratigraphy, owing to the layers of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1362\"><strong>volcanic tufts<\/strong> <\/a> that have accumulated over millions of years. <span style=\"background-color: #ff9900\">These tufts may be dated using absolute dating techniques, such as Potassium-Argon dating (described in Chapter 7). This means that it is possible to know a relatively refined date for any fossil if the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1364\"><strong>context<\/strong> <\/a> (i.e., exact location) of that find is known.<\/span> Similarly, comparisons between the faunal assemblages of these stratigraphic layers have allowed researchers to chronologically identify environmental changes.<\/span><\/p>\n<figure style=\"width: 313px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image20-1-1.png\" alt=\"Occlusal view of an Au. anamensis mandible, with relatively large teeth, including canines.\" width=\"313\" height=\"313\" \/><figcaption class=\"wp-caption-text\">Figure 9.13: As seen in this mandible of KNM-KP 29281, Australopithecus anamensis had relatively large canine teeth. Credit: <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20anamensis\/KNM-KP%2029281\">Australopithecus anamensis: KNM-KP 29281 occlusal view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p><span style=\"color: #000000\">The earliest known Australopithecine is dated to 4.2 mya to 3.8 mya. <strong><em>Australopithecus anamensis<\/em><\/strong> (after \u201cAnam,\u201d meaning \u201clake\u201d from the Turkana region in Kenya; Leakey et al. 1995; Patterson and Howells 1967) is currently found from sites in the Turkana region (Kenya) and Middle Awash (Ethiopia; Figure 9.13). Recently, a 2019 find from Ethiopia, named MRD, after Miro Dora where it was found, was discovered by an Ethiopian herder named Ali Bereino. It is one of the most complete cranial finds of this species (Ward et al. 1999). A small brain size (370 cc), relatively large canines, projecting cheekbones, and earholes show more ancestral features as compared to those of more recent Australopithecines. The most important element discovered with this species is a fragment of a tibia (shinbone), which demonstrates features associated with weight transfer during bipedal walking. Similarly, the earliest found hominin femur belongs to this species. Ancestral traits in the upper limb (such as the humerus) indicate some retained arboreal locomotion.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Some researchers suggest that <em>Au. anamensis<\/em> is an intermediate form of the chronospecies that becomes <em>Au. afarensis<\/em>, evolving from <em>Ar. ramidus<\/em>. However, this is debated, with other researchers suggesting morphological similarities and affinities with more recent species instead. Almost 100 specimens, representing over 20 individuals, have been found to date (Leakey et al. 1995; McHenry 2009; Ward et al. 1999).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Au. afarensis<\/em><\/strong> is one of the oldest and most well-known australopithecine species and consists of a large number of fossil remains. <em>Au. afarensis<\/em> (which means \u201cfrom the Afar region\u201d) is dated to between 2.9 mya and 3.9 mya and is found in sites all along the EARS system, in Tanzania, Kenya, and Ethiopia (Figure 9.14). The most famous individual from this species is a partial female skeleton discovered in Hadar (Ethiopia), later nicknamed \u201cLucy,\u201d after the Beatles\u2019 song \u201cLucy in the Sky with Diamonds,\u201d which was played in celebration of the find (Johanson et al. 1978; Kimbel and Delezene 2009). This skeleton was found in 1974 by Donald Johanson and dates to approximately 3.2 mya. In addition, in 2002 a juvenile of the species was found by Zeresenay Alemseged and given the name \u201cSelam\u201d (meaning \u201cpeace,\u201d DIK 1-1), though it is popularly known as \u201cLucy\u2019s Child\u201d or as the \u201cDikika Child\u201d (Alemseged et al. 2006). Similarly, the \u201cLaetoli Footprints\u201d (discussed in Chapter 7; Hay and Leakey 1982; Leakey and Hay 1979) have drawn much attention.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 643px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-295 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.14.jpg\" alt=\"Two images of life-like reconstructions of female and male Au. afarensis.\" width=\"643\" height=\"322\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.14 a-b: Artistic reconstructions of Australopithecus afarensis by artist John Gurche. Female \u201cLucy\u201d is left and a male is on the right. Credit: a. <a class=\"rId106\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Australopithecus afarensis, \u201cLucy,\u201d adult female. Reconstruction based on AL-288-1 by artist John Gurche, front view close-up<\/a> by <a class=\"rId107\" href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?\u201d] is <a class=\"rId108\" href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>; b. <a class=\"rId109\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Australopithecus afarensis, adult male. Reconstruction based on <\/a><a class=\"rId110\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">AL444-2<\/a><a class=\"rId111\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\"> by John Gurche<\/a> by <a class=\"rId112\" href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?\u201d] is <a class=\"rId113\" href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>.<\/figcaption><\/figure>\n<figure style=\"width: 320px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image40.png\" alt=\"A partial skeleton includes bones of the cranium, mandible, and postcranium.\" width=\"320\" height=\"772\" \/><figcaption class=\"wp-caption-text\">Figure 9.15: The humanlike femoral neck, valgus knee, and bowl-shaped hip seen in the \u201cLucy\u201d skeleton indicates that Australopithecus afarensis was bipedal. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Lucy_blackbg.jpg\">Lucy blackbg<\/a> [AL 288-1, Australopithecus afarensis, cast from Museum national d'histoire naturelle, Paris] by 120 is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The canines and molars of <em>Au. afarensis<\/em> are reduced relative to great apes but are larger than those found in modern humans (indicative of a generalist diet); in addition, <em>Au. afarensis <\/em>has a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1366\"><strong>prognathic<\/strong> <\/a> face (the face below the eyes juts anteriorly) and robust facial features that indicate relatively strong chewing musculature (compared with <em>Homo<\/em>) but which are less extreme than in <em>Paranthropus<\/em>. Despite a reduction in canine size in this species, large overall size variation indicates high levels of sexual dimorphism.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Skeletal evidence indicates that this species was bipedal, as its pelvis and lower limb demonstrate a humanlike femoral neck, valgus knee, and bowl-shaped hip (Figure 9.15). <span style=\"background-color: #ccffcc\">More evidence of bipedalism is found in the footprints of this species<\/span>. <em>Au. afarensis<\/em> is associated with the Laetoli Footprints, <span style=\"background-color: #ff9900\">a 24-meter trackway of hominin fossil footprints preserved in volcanic ash discovered by Mary Leakey in Tanzania and dated to 3.5 mya to 3 mya. This set of prints is thought to have been produced by three bipedal individuals as there are no knuckle imprints, no opposable big toes, and a clear arch is present. The infants of this species are thought to have been more arboreal than the adults, as discovered through analyses of the foot bones of the Dikika Child dated to 3.32 mya (Alemseged et al. 2006).<\/span><\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Although not found in direct association with stone tools, potential evidence for cut marks on bones, found at Dikika, and dated to 3.39 mya indicates a possible temporal\/ geographic overlap between meat eating, tool use, and this species. However, this evidence is fiercely debated. Others have associated the cut marks with the earliest tool finds from Lomekwi, Kenya, temporally (3.3 mya) and in close geographic proximity to this species.<\/span><\/p>\n<h4 class=\"import-Normal\"><em><span style=\"color: #000000\">South African Australopithecines<\/span><\/em><\/h4>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Since the discovery of the Taung Child, there have been numerous Australopithecine discoveries from the region known as \u201cThe Cradle of Humankind,\u201d which was recently given UNESCO World Heritage Site status as \u201cThe Fossil Hominid Sites of South Africa.\u201d The limestone caves found in the Cradle allow for the excellent preservation of fossils. Past animals navigating the landscape and falling into cave openings, or caves used as dens by carnivores, led to the accumulation of deposits over millions of years. Many of the hominin fossils, encased in <strong>breccia<\/strong> (hard, calcareous sedimentary rock), are recently exposed from limestone quarries mined in the previous century. This means that extracting fossils requires excellent and detailed exposed work, often by a team of skilled technicians.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">While these sites have historically been difficult to date, with mixed assemblages accumulated over large time periods, advances in techniques such as uranium-series dating have allowed for greater accuracy. Historically, the excellent faunal record from East Africa has been used to compare sites based on <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1368\"><strong>relative dating<\/strong>,<\/a> whereby environmental and faunal changes and extinction events allow us to know which hominin finds are relatively younger or older than others.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The discovery of the Taung Child in 1924 (discussed in the Special Topic box \u201cThe Taung Child\u201d below) shifted the focus of palaeoanthropological research from Europe to Africa, although acceptance of this shift was slow (Broom 1947; Dart 1925). The species to which it is assigned, <strong><em>Australopithecus africanus<\/em><\/strong> (name meaning \u201cSouthern Ape of Africa\u201d), is currently dated to between 3.3 mya and 2.1 mya (Pickering and Kramers 2010), with discoveries from Sterkfontein, Taung, Makapansgat, and Gladysvale in South Africa (Figure 9.16). A relatively large brain (400 cc to 500 cc), small canines without an associated diastema, and more rounded cranium and smaller teeth than <em>Au. afarensis<\/em> indicate some derived traits. Similarly, the postcranial remains (in particular, the pelvis) indicate bipedalism. However, the sloping face and curved phalanges (indicative of retained arboreal locomotor abilities) show some ancestral features. Although not in direct association with stone tools, a 2015 study noted that the trabecular bone morphology of the hand was consistent with forceful tool manufacture and use, suggesting potential early tool abilities.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 570px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image19-1.png\" alt=\"A life-like reconstruction of the face of Australopithecus africanus, smiling in anterior view.\" width=\"570\" height=\"570\" \/><figcaption class=\"wp-caption-text\">Figure 9.16: An artistic reconstruction of Australopithecus africanus by John Gurche. Credit: <a href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Australopithecus africanus. Reconstruction based on STS 5 by John Gurche <\/a>by <a href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?] is <a href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Another famous <em>Au. africanus<\/em> skull (the skull of \u201cMrs. Ples\u201d) was previously attributed to <em>Plesianthropus transvaalensis<\/em><em>, <\/em>meaning \u201cnear human from the Transvaal,\u201d the old name for Gauteng Province, South Africa (Broom 1947, 1950). The name was shortened by contemporary journalists to \u201cPles\u201d (Figure 9.17). Due to the prevailing mores of the time, the assumed female found herself married, at least in name, and has become widely known as \u201cMrs. Ples.\u201d It was later reassigned to <em>Au. africanus<\/em> and is now argued by some to be a young male rather than an adult female cranium (Thackeray 2000, Thackeray et al. 2002).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 548px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-298 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.17.jpg\" alt=\"Four views of an ancient skull are shown on a black background.\" width=\"548\" height=\"548\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.17: The \u201cMrs. Ples\u201d brain case is small in size (like apes) but its face is less prognathic; its foramen magnum is positioned more like a modern human than an African apes. Credit: a. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 anterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 posterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 superior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; and d. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 lateral right view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In 2008, nine-year-old Matthew Berger, son of paleoanthropologist Lee Berger, noted a clavicle bone in some leftover mining breccia in the Malapa Fossil Site (South Africa). After rigorous studies, the species, <strong><em>Australopithecus sediba<\/em><\/strong> (meaning \u201cfountain\u201d or \u201cwellspring\u201d in the South African language of Sesotho), was named in 2010 (Figure 9.18; Berger et al. 2010). The first type specimen belongs to a juvenile male, Karabo (MH1), but the species is known from at least six partial skeletons, from infants through adults. These specimens are currently dated to 1.97 mya (Dirks et al. 2010). The discoverers have argued that <em>Au. sediba<\/em> shows mosaic features between <em>Au. africanus<\/em> and the genus, <em>Homo<\/em>, which potentially indicates a transitional species, although this is heavily debated. These features include a small brain size (<em>Australopithecus<\/em>-like; 420 cc to 450 cc) but gracile mandible and small teeth (<em>Homo<\/em>-like). Similarly, the postcranial skeletons are also said to have mosaic features: scientists have interpreted this mixture of traits (such as a robust ankle but evidence for an arch in the foot) as a transitional phase between a body previously adapted to arborealism (particularly in evidence from the bones of the wrist) to one that adapted to bipedal ground walking. Some researchers have argued that <em>Au. sediba<\/em> shows a modern hand morphology (shorter fingers and a longer thumb), indicating that adaptations to tool manufacture and use may be present in this species.<\/span><\/p>\n<figure style=\"width: 531px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image17-1.png\" alt=\"A beige-colored skull with no mandible on a black background has some missing teeth.\" width=\"531\" height=\"400\" \/><figcaption class=\"wp-caption-text\">Figure 9.18: Australopithecus sediba shows mosaic features between Au. africanus and Homo. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Australopithecus_sediba.JPG\">Australopithecus sediba<\/a>, photo by Brett Eloff courtesy <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Profberger\">Profberger<\/a> and <a href=\"https:\/\/en.wikipedia.org\/wiki\/University_of_the_Witwatersrand\">Wits University<\/a>, is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Another famous Australopithecine find from South Africa is that of the nearly complete skeleton now known as \u201cLittle Foot\u201d (Clarke 1998, 2013). Little Foot (StW 573) is potentially the earliest dated South African hominin fossil, dating to 3.7 mya, based on radiostopic techniques, although some argue that it is younger than 3 mya (Pickering and Kramers 2010). The name is jokingly in contrast to the cryptid species \u201cbigfoot\u201d and is named because the initial discovery of four ankle bones indicated bipedality. Little Foot was discovered by Ron Clarke in 1994, when he came across the ankle bones while sorting through monkey fossils in the University of Witwatersrand collections (Clarke and Tobias 1995). He asked Stephen Motsumi and Nkwane Molefe to identify the known records of the fossils, which allowed them to find the rest of the specimen within just days of searching the Sterkfontein Caves\u2019 Silberberg Grotto.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The discoverers of Little Foot insist that other fossil finds, previously identified as <em>Au. Africanus<\/em>, be placed in this new species based on shared ancestral traits with older East African Australopithecines (Clarke and Kuman 2019). These include features such as a relatively large brain size (408 cc), robust zygomatic arch, and a flatter midface. Furthermore, the discoverers have argued that the heavy anterior dental wear patterns, relatively large anterior dentition, and smaller hind dentition of this specimen more closely resemble that of <em>Au. anamensis<\/em> or <em>Au. afarensis<\/em>. It has thus been placed in the species <strong><em>Australopithecus prometheus<\/em><\/strong>. This species name refers to a previously defunct taxon named by Raymond Dart. The species designation was, through analyzing Little Foot, revived by Ron Clarke, who insists that many other fossil hominin specimens have prematurely been placed into <em>Au. africanus<\/em>. Others say that it is more likely that <em>Au. africanus<\/em> is a more variable species and not representative of two distinct species.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\"><em>Paranthropus<\/em> \u201cRobust\u201d Australopithecines<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In the robust australopithecines, the specialized nature of the teeth and masticatory system, such as flaring zygomatic arches (cheekbones), accommodate very large temporalis (chewing) muscles. These features also include a large, broad, dish-shaped face and and a large mandible with extremely large posterior dentition (referred to as megadonts) and hyper-thick enamel (Kimbel 2015; Lee-Thorp 2011; Wood 2010). Research has revolved around the shared adaptations of these \u201crobust\u201d australopithecines, linking their morphologies to a diet of hard and\/or tough foods (Brain 1967; Rak 1988). Some argued that the diet of the robust australopithecines was so specific that any change in environment would have accelerated their extinction. The generalist nature of the teeth of the gracile australopithecines, and of early <em>Homo<\/em>, would have made them more capable of adapting to environmental change. However, some have suggested that the features of the robust australopithecines might have developed as an effective response to what are known as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1370\"><strong>fallback <\/strong><strong>foods<\/strong><\/a> in hard times rather than indicating a lack of adaptability.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are currently three widely accepted robust australopithecus or, <em>Paranthropus<\/em>, species: <em>P. aethiopic<\/em><em>us<\/em>, which has more ancestral traits, and <em>P. boisei and P. robustus<\/em>, which are more derived in their features (Strait et al. 1997; Wood and Schroer 2017). These three species have been grouped together by a majority of scholars as a single genus as they share more derived features (are more closely related to each other; or, in other words, are <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1372\"><strong>monophyletic<\/strong><\/a>) than the other australopithecines (Grine 1988; Hlazo 2015; Strait et al. 1997; Wood 2010 ). While researchers have mostly agreed to use the umbrella term <em>Paranthropus<\/em>, there are those who disagree (Constantino and Wood 2004, 2007; Wood 2010).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">As a collective, this genus spans 2.7 mya to 1.0 mya, although the dates of the individual species differ. The earliest of the Paranthropus species, <strong><em>Paranthropus aethiopicus<\/em><\/strong>, is dated to between 2.7 mya and 2.3 mya and currently found in Tanzania, Kenya, and Ethiopia in the EARS system (Figure 9.19; Constantino and Wood 2007; Hlazo 2015; Kimbel 2015; Walker et al. 1986; White 1988). It is well known because of one specimen known as the \u201cBlack Skull\u201d (KNM\u2013WT 17000), so called because of the mineral manganese that stained it black during fossilization (Kimbel 2015). As with all robust Australopithecines, <em>P. aethiopicus<\/em> has the shared derived traits of large, flat premolars and molars; large, flaring zygomatic arches for accommodating large chewing muscles (the temporalis muscle); a sagittal crest (ridge on the top of the skull) for increased muscle attachment of the chewing muscles to the skull; and a robust mandible and supraorbital torus (brow ridge). However, only a few teeth have been found. A proximal tibia indicates bipedality and similar body size to <em>Au. afarensis<\/em>. In recent years, researchers have discovered and assigned a proximal tibia and juvenile cranium (L.338y-6) to the species (Wood and Boyle 2016).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 666px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-300 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.19.jpg\" alt=\"Five views of a beige partial skull on a black background.\" width=\"666\" height=\"444\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.19: The \u201cBlack Skull\u201d (Paranthropus aethiopicus) had a large sagittal crest and large, flared zygomatic arches that indicate it had large chewing muscles and a powerful biting force. Credit: a. <a class=\"rId156\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId157\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 anterior view<\/a> by \u00a9<a class=\"rId158\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId159\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId160\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId161\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId162\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 lateral right view<\/a> by \u00a9<a class=\"rId163\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId164\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId165\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a class=\"rId166\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId167\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 superior view<\/a> by \u00a9<a class=\"rId168\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId169\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId170\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; d. <a class=\"rId171\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId172\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 posterior view<\/a> by \u00a9<a class=\"rId173\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId174\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId175\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; e. <a class=\"rId176\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId177\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 inferior view<\/a> by \u00a9<a class=\"rId178\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId179\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId180\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">First attributed as <em>Zinjanthropus boisei<\/em> (with the first discovery going by the nickname \u201cZinj\u201d or sometimes \u201cNutcracker Man\u201d), <strong><em>Paranthropus boisei<\/em><\/strong> was discovered in 1959 by Mary Leakey (see Figure 9.20 and 9.21; Hay 1990; Leakey 1959). This \u201crobust\u201d australopith species is distributed across countries in East Africa at sites such as Kenya (Koobi Fora, West Turkana, and Chesowanja), Malawi (Malema-Chiwondo), Tanzania (Olduvai Gorge and Peninj), and Ethiopia (Omo River Basin and Konso). The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1374\">hypodigm<\/a><\/strong>, sample of fossils whose features define the group, has been found by researchers to date to roughly 2.4 mya to 1.4 mya. Due to the nature of its exaggerated, larger, and more robust features, <em>P. boisei <\/em>has been termed <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1376\"><strong>hyper-robust<\/strong><\/a>\u2014that is, even more heavily built than other robust species, with very large, flat posterior dentition (Kimbel 2015). Tools dated to 2.5 mya in Ethiopia have been argued to possibly belong to this species. Despite the cranial features of <em>P. boisei<\/em> indicating a tough diet of tubers, nuts, and seeds, isotopes indicate a diet high in C4 foods (e.g., grasses, such as sedges). Another famous specimen from this species is the Peninj mandible from Tanzania, found in 1964 by Kimoya Kimeu.<\/span><\/p>\n<figure style=\"width: 557px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image14-1.png\" alt=\"Life-like reconstruction of Paranthropus boisei.\" width=\"557\" height=\"557\" \/><figcaption class=\"wp-caption-text\">Figure 9.20: Artistic reconstruction of a Paranthropus boisei, male, by John Gurche. Credit: <a href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Paranthropus boisei, male. Reconstruction based on OH 5 and KNM-ER 406 by John Gurche<\/a> by <a href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?\u201d] is <a href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>.<\/figcaption><\/figure>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 565px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-302 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.21.jpg\" alt=\"Three views of an ancient skull are shown on a black background.\" width=\"565\" height=\"565\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.21: \u201cNutcracker Man\u201d (Paranthropus boisei) had hyper-robust features including very large dentition, flaring zygomatic arches, a broad concave face. It had a powerful and extremely efficient chewing force. Credit: <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20boisei\/OH%205\">Paranthropus boisei: OH 5 anterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20boisei\/OH%205\">Paranthropus boisei: OH 5 inferior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20boisei\/OH%205\">Paranthropus boisei: OH 5 posterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Paranthropus robustus<\/em><\/strong> was the first taxon to be discovered within the genus in Kromdraai B by a schoolboy named Gert Terblanche; subsequent fossil discoveries were made by researcher Robert Broom in 1938 (Figure 9.22; Broom 1938a, 1938b, 1950), with the holotype specimen TM 1517 (Broom 1938a, 1938b, 1950; Hlazo 2018). <em>Paranthropus robustus<\/em> dates approximately from 2.0 mya to 1 mya and is the only taxon from the genus to be discovered in South Africa. Several of these fossils are fragmentary in nature, distorted, and not well preserved because they have been recovered from quarry breccia using explosives. <em>P. robustus<\/em> features are neither as \u201chyper-robust\u201d as <em>P. boisei<\/em> nor as ancestral as <em>P. aethiopicus<\/em>; instead, they have been described as being less derived, more general features that are shared with both East African species (e.g., the sagittal crest and zygomatic flaring; Rak 1983; Walker and Leakey 1988). Enamel hypoplasia is also common in this species, possibly because of instability in the development of large, thick-enameled dentition.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 572px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-303 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.22.jpg\" alt=\"Four views of a beige-colored skull are shown on a black background.\" width=\"572\" height=\"619\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.22: SK 48, a Paranthropus robustus specimen, had less derived, more general features that were not as robust as P. boisei and not as ancestral as P. aethiopicus. Credit: a. <a class=\"rId208\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId209\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 anterior view<\/a> by \u00a9<a class=\"rId210\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId211\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId212\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId213\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId214\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 superior view<\/a> by \u00a9<a class=\"rId215\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId216\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId217\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a class=\"rId218\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId219\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 inferior view<\/a> by \u00a9<a class=\"rId220\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId221\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId222\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; d. <a class=\"rId223\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId224\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 lateral left view<\/a> by \u00a9<a class=\"rId225\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId226\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId227\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Comparisons between Gracile and Robust Australopiths<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Comparisons between gracile and robust australopithecines may indicate different phylogenetic groupings or parallel evolution in several species. In general, the robust australopithecines have large temporalis (chewing) muscles, as indicated by flaring zygomatic arches, sagittal crests, and robust mandibles (jawbones). Their hind dentition is large (megadont), with low cusps and thick enamel. Within the gracile australopithecines, researchers have debated the relatedness of the species, or even whether these species should be lumped together to represent more variable or polytypic species. Often researchers will attempt to draw chronospecific trajectories, with one taxon said to evolve into another over time.<\/span><\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Special Topic: The Taung Child<\/span><\/h2>\n<figure id=\"attachment_304\" aria-describedby=\"caption-attachment-304\" style=\"width: 570px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-304 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.23.jpg\" alt=\"An ancient skull in anterior and lateral views. One view shows an imprint of the brain.\" width=\"570\" height=\"285\" \/><figcaption id=\"caption-attachment-304\" class=\"wp-caption-text\">Figure 9.23: The Taung Child has a nearly complete face, mandible, and partial endocranial cast. Credit: a. <em>A<\/em><a class=\"rId230\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\"><em>ustralopithecus africanus<\/em><\/a><a class=\"rId231\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\">: Taung 1 anterior view<\/a> by \u00a9<a class=\"rId232\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId233\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId234\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId235\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\"><em>australopithecus africanus<\/em><\/a><a class=\"rId236\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\">: Taung 1 lateral right view<\/a> by \u00a9<a class=\"rId237\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId238\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId239\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p>&nbsp;<\/p>\n<p><span style=\"color: #000000\">The well-known fossil of a juvenile <em>Australopithecine<\/em>, the \u201cTaung Child,\u201d was the first early hominin evidence ever discovered and was the first to demonstrate our common human heritage in Africa (Figure 9.23; Dart 1925). The tiny facial skeleton and natural endocast were discovered in 1924 by a local quarryman in the North West Province in South Africa and were painstakingly removed from the surrounding cement-like breccia by Raymond Dart using his wife\u2019s knitting needles. When first shared with the scientific community in 1925, it was discounted as being nothing more than a young monkey of some kind. Prevailing biases of the time made it too difficult to contemplate that this small-brained hominin could have anything to do with our own history. The fact that it was discovered in Africa simply served to strengthen this bias.<\/span><\/p>\n<\/div>\n<p>&nbsp;<\/p>\n<h2><span style=\"color: #000000\">Early Tool Use and Technology<br \/>\n<\/span><\/h2>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Early Stone Age Technology (ESA)<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1378\">Early Stone Age (ESA)<\/a><\/strong> marks the beginning of recognizable technology made by our human ancestors. Stone-tool (or <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1380\"><strong>lithic<\/strong><\/a>) technology is defined by the fracturing of rocks and the manufacture of tools through a process called <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1382\"> <strong>knapping<\/strong><\/a>. The Stone Age lasted for more than 3 million years and is broken up into chronological periods called the Early (ESA), Middle (MSA), and Later Stone Ages (LSA). Each period is further broken up into a different <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1384\">techno-complex<\/a><\/strong>, a term encompassing multiple <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1386\">assemblages<\/a><\/strong> (collections of artifacts) that share similar traits in terms of artifact production and morphology. The ESA spanned the largest technological time period of human innovation from over 3 million years ago to around 300,000 years ago and is associated almost entirely with hominin species prior to modern <em>Homo sapiens. <\/em>As the ESA advanced, stone tool makers (known as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1390\"><strong>knappers<\/strong><\/a>) began to change the ways they detached <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1388\">flakes<\/a><\/strong> and eventually were able to shape artifacts into functional tools. These advances in technology go together with the developments in human evolution and cognition, dispersal of populations across the African continent and the world, and climatic changes.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In order to understand the ESA, it is important to consider that not all assemblages are exactly the same within each techno-complex: one can have multiple phases and traditions at different sites (Lombard et al. 2012). However, there is an overarching commonality between them. Within stone tool assemblages, both flakes or <strong>cores<\/strong> (the rocks from which flakes are removed) are used as tools.<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1392\"> <strong>Large Cutting Tools (LCTs)<\/strong><\/a> are tools that are shaped to have functional edges. It is important to note that the information presented here is a small fraction of what is known about the ESA, and there are ongoing debates and discoveries within archaeology.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Currently, the oldest-known stone tools, which form the techno-complex the Lomekwian, date to 3.3 mya (Harmand et al. 2015; Toth 1985). They were found at a site called Lomekwi 3 in Kenya. This techno-complex is the most recently defined and pushed back the oldest-known date for lithic technology. There is only one known site thus far and, due to the age of the site, it is associated with species prior to <em>Homo<\/em>, such as <em>Kenyanthropus platyops.<\/em> Flakes were produced through indirect percussion, whereby the knappers held a rock and hit it against another rock resting on the ground. The pieces are very chunky and do not display the same fracture patterns seen in later techno-complexes. Lomekwian knappers likely aimed to get a sharp-edged piece on a flake, which would have been functional, although the specific function is currently unknown.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Stone tool use, however, is not only understood through the direct discovery of the tools. Cut marks on fossilized animal bones may illuminate the functionality of stone tools. In one controversial study in 2010, researchers argued that cut marks on a pair of animal bones from Dikika (Ethiopia), dated to 3.4 mya, were from stone tools. The discoverers suggested that they be more securely associated, temporally, with <em>Au. afarensis<\/em>. However, others have noted that these marks are consistent with teeth marks from crocodiles and other carnivores.<\/span><\/p>\n<figure style=\"width: 324px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image29-1.png\" alt=\"A technical line drawing of an Oldowan chopper.\" width=\"324\" height=\"275\" \/><figcaption class=\"wp-caption-text\">Figure 9.24: Some scholars believe that some genera explored in this chapter were capable of producing more complex stone tools (Oldowan). Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Olduwan_Industry_Chopper_2.jpeg\">Olduwan Industry Chopper 2<\/a> by Emmyanne29 is under a <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/legalcode\">CC0 1.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The Oldowan techno-complex is far more established in the scientific literature (Leakey 1971). It is called the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1394\">Oldowan<\/a><\/strong> because it was originally discovered in Olduvai Gorge, Tanzania, but the oldest assemblage is from Gona in Ethiopia, dated to 2.6 mya (Semaw 2000). The techno-complex is defined as a core and flake industry. Like the Lomekwian, there was an aim to get sharp-edged flakes, but this was achieved through a different production method. Knappers were able to actively hold or manipulate the core being knapped, which they could directly hit using a hammerstone. This technique is known as free-hand percussion, and it demonstrates an understanding of fracture mechanics. It has long been argued that the Oldowan hominins were skillful in tool manufacture.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Because Oldowan knapping requires skill, earlier researchers have attributed these tools to members of our genus, <em>Homo<\/em>. However, some have argued that these tools are in more direct association with hominins in the genera described in this chapter (Figure 9.24).<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Invisible Tool Manufacture and Use<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The vast majority of our understanding of these early hominins comes from fossils and reconstructed paleoenvironments. It is only from 3 mya when we can start \u201clooking into their minds\u201d and lifestyles by analyzing their manufacture and use of stone tools. However, the vast majority of tool use in primates (and, one can argue, in humans) is not with durable materials like stone. All of our extant great ape relatives have been observed using sticks, leaves, and other materials for some secondary purpose (to wade across rivers, to \u201cfish\u201d for termites, or to absorb water for drinking). It is possible that the majority of early hominin tool use and manufacture may be invisible to us because of this preservation bias.<\/span><\/p>\n<h2 class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000;background-color: #ff00ff\">Chapter Summary<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000;background-color: #ff00ff\">The fossil record of our earliest hominin relatives has allowed paleoanthropologists to unpack some of the mysteries of our evolution. We now know that traits associated with bipedalism evolved before other \u201chuman-like\u201d traits, even though the first hominins were still very capable of arboreal locomotion. We also know that, for much of this time, hominin taxa were diverse in the way they looked and what they ate, and they were widely distributed across the African continent. And we know that the environments in which these hominins lived underwent many changes over this time during several warming and cooling phases.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;background-color: #ff00ff\">Yet this knowledge has opened up many new mysteries. We still need to better differentiate some taxa. In addition, there are ongoing debates about why certain traits evolved and what they meant for the extinction of some of our relatives (like the robust australopiths). The capabilities of these early hominins with respect to tool use and manufacture is also still uncertain.<\/span><\/p>\n<h2 class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Hominin Species Summaries<br \/>\n<\/span><\/h2>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Sahelanthropus tchadensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">7 mya to 6 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Chad<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The initial discovery, made in 2001.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">360 cc average<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Smaller than in extant great apes; larger and pointier than in humans. Canines worn at the tips.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A short cranial base and a foramen magnum (hole in which the spinal cord enters the cranium) that is more humanlike in positioning; has been argued to indicate upright walking.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Currently little published postcranial material.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The extent to which this hominin was bipedal is currently heavily debated. If so, it would indicate an arboreal bipedal ancestor of hominins, not a knuckle-walker like chimpanzees.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Orrorin tugenensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">6 mya to 5.7 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Tugen Hills (Kenya)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Original discovery in 2000.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Smaller cheek teeth (molars and premolars) than even more recent hominins (i.e., derived), thick enamel, and reduced, but apelike, canines.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Not many found<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Fragmentary leg, arm, and finger bones have been found. Indicates bipedal locomotion.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Potential toolmaking capability based on hand morphology, but nothing found directly.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">This is the earliest species that clearly indicates adaptations for bipedal locomotion.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Ardipithecus kadabba<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">5.2 mya to 5.8 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Middle Awash (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Discovered by Yohannes Haile-Selassie in 1997.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Larger hind dentition than in modern chimpanzees. Thick enamel and larger canines than in later hominins.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A large hallux (big toe) bone indicates a bipedal \u201cpush off.\u201d<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Faunal evidence indicates a mixed grassland\/woodland environment.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Ardipithecus ramidus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">4.4 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Middle Awash region and Gona (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">A partial female skeleton nicknamed \u201cArdi\u201d (ARA-VP-6\/500) (found in 1994).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">300 cc to 350 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Little differences between the canines of males and females (small sexual dimorphism).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Midfacial projection, slightly prognathic. Cheekbones less flared and robust than in later hominins.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Ardi demonstrates a mosaic of ancestral and derived characteristics in the postcrania. For instance, an opposable big toe similar to chimpanzees (i.e., more ancestral), which could have aided in climbing trees effectively. However, the pelvis and hip show that she could walk upright (i.e., it is derived), supporting her hominin status.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">None directly associated<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">Over 110 specimens from Aramis<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus anamensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">4.2 mya to 3.8 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Turkana region (Kenya); Middle Awash (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A 2019 find from Ethiopia, named MRD.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">370 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Relatively large canines compared with more recent Australopithecines.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Projecting cheekbones and ancestral earholes.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lower limb bones (tibia and femur) indicate bipedality; arboreal features in upper limb bones (humerus) found.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Almost 100 specimens, representing over 20 individuals, have been found to date.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus afarensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">3.9 mya to 2.9 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Afar Region, Omo, Maka, Fejej, and Belohdelie (Ethiopia); Laetoli (Tanzania); Koobi Fora (Kenya)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lucy (discovery: 1974), Selam (Dikika Child, discovery: 2000), Laetoli Footprints (discovery: 1976).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">380 cc to 430 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Reduced canines and molars relative to great apes but larger than in modern humans.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Prognathic face, facial features indicate relatively strong chewing musculature (compared with <em>Homo<\/em>) but less extreme than in <em>Paranthropus<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clear evidence for bipedalism from lower limb postcranial bones. Laetoli Footprints indicate humanlike walking. Dikika Child bones indicate retained ancestral arboreal traits in the postcrania.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">None directly, but close in age and proximity to controversial cut marks at Dikika and early tools in Lomekwi.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Au. afarensis<\/em> is one of the oldest and most well-known australopithecine species and consists of a large number of fossil remains.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus bahrelghazali<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">3.6 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Chad<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">\u201cAbel,\u201d the holotype (discovery: 1995).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Arguably within range of variation of <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus prometheus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">3.7 mya (debated)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">Sterkfontein (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">\u201cLittle Foot\u201d (StW 573) (discovery: 1994)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">408 cc (Little Foot estimate)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">Heavy anterior dental wear patterns, relatively large anterior dentition and smaller hind dentition, similar to <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">Relatively larger brain size, robust zygomatic arch, and a flatter midface.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">The initial discovery of four ankle bones indicated bipedality.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">Highly debated new species designation.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus <\/em><em>deyiremada<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">3.5 mya to 3.3 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Woranso-Mille (Afar region, Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">First fossil mandible bones were discovered in 2011 in the Afar region of Ethiopia by Yohannes Haile-Selassie.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Smaller teeth with thicker enamel than seen in <em>Au. afarensis<\/em>, with a potentially hardier diet.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Larger mandible and more projecting cheekbones than in <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Contested species designation; arguably a member of <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Kenyanthopus<\/em><em> platyops<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">3.5 mya to 3.2 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Lake Turkana (Kenya)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">KNM\u2013WT 40000 (discovered 1999)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Difficult to determine but appears within the range of <em>Australopithecus afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Small molars\/dentition (<em>Homo<\/em>-like characteristic)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Flatter (i.e., orthognathic) face<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Some have associated the earliest tool finds from Lomekwi, Kenya, temporally (3.3 mya) and in close geographic proximity to this species\/specimen.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Taxonomic placing of this species is quite divided. The discoverers have argued that this species is ancestral to <em>Homo<\/em>, in particular to <em>Homo <\/em><em>ruldolfensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Australopithecus africanus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">3.3 mya to 2.1 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Sterkfontein, Taung, Makapansgat, Gladysvale (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Taung Child (discovery in 1994), \u201cMrs. Ples\u201d (discover in 1947), Little Foot (arguable; discovery in 1994).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">400 cc to 500 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Smaller teeth (derived) relative to <em>Au. afarensis<\/em>. Small canines with no diastema.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">A rounder skull compared with <em>Au. afarensis<\/em> in East Africa. A sloping face (ancestral).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Similar postcranial evidence for bipedal locomotion (derived pelvis) with retained arboreal locomotion, e.g., curved phalanges (fingers), as seen in <em>Au. afarensis.<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">None with direct evidence.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">A 2015 study noted that the trabecular bone morphology of the hand was consistent with forceful tool manufacture and use, suggesting potential early tool abilities.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus garhi<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.5 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Middle Awash (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">450 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Larger hind dentition than seen in other gracile Australopithecines.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A femur of a fragmentary partial skeleton, argued to belong to <em>Au. garhi<\/em>, indicates this species may be longer-limbed than <em>Au. afarensis<\/em>, although still able to move arboreally.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Crude stone tools resembling Oldowan (described later) have been found in association with <em>Au. garhi<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">This species is not well documented or understood and is based on only a few fossil specimens.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Paranthropus aethiopicus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.7 mya to 2.3 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">West Turkana (Kenya); Laetoli (Tanzania); Omo River Basin (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The \u201cBlack Skull\u201d (KNM\u2013WT 17000) (discovery 1985).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain Size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">410 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>P. aethiopicus<\/em> has the shared derived traits of large flat premolars and molars, although few teeth have been found.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Large flaring zygomatic arches for accommodating large chewing muscles (the temporalis muscle), a sagittal crest for increased muscle attachment of the chewing muscles to the skull, and a robust mandible and supraorbital torus (brow ridge).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A proximal tibia indicates bipedality and similar size to <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The \u201cBlack Skull\u201d is so called because of the mineral manganese that stained it black during fossilization.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Paranthropus boisei<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.4 mya to 1.4 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Koobi Fora, West Turkana, and Chesowanja (Kenya); Malema-Chiwondo (Malawi), Olduvai Gorge and Peninj (Tanzania); and Omo River basin and Konso (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">\u201cZinj,\u201d or sometimes \u201cNutcracker Man\u201d (OH5), in 1959 by Mary Leakey. The Peninj mandible from Tanzania, found in 1964 by Kimoya Kimeu.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">500 cc to 550 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Very large, flat posterior dentition (largest of all hominins currently known). Much smaller anterior dentition. Very thick dental enamel.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Indications of very large chewing muscles (e.g., flaring zygomatic arches and a large sagittal crest).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Evidence for high variability and sexual dimorphism, with estimates of males at 1.37 meters tall and females at 1.24 meters.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Richard Leakey and Bernard Wood have both suggested that<em> P. boisei<\/em> could have made and used stone tools. Tools dated to 2.5 mya in Ethiopia have been argued to possibly belong to this species.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Despite the cranial features of <em>P. boisei<\/em> indicating a tough diet of tubers, nuts, and seeds, isotopes indicate a diet high in C4 foods (e.g., grasses, such as sedges). This differs from what is seen in<em> P. robustus<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Australopithecus sediba<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">1.97 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Malapa Fossil Site (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Karabo (MH1) (discovery in 2008)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">420 cc to 450 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Small dentition with Australopithecine cusp-spacing.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Small brain size (<em>Australopithecus<\/em>-like) but gracile mandible (<em>Homo<\/em>-like).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Scientists have interpreted this mixture of traits (such as a robust ankle but evidence for an arch in the foot) as a transitional phase between a body previously adapted to arborealism (tree climbing, particularly in evidence from the bones of the wrist) to one that adapted to bipedal ground walking.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">None of direct association, but some have argued that a modern hand morphology (shorter fingers and a longer thumb) means that adaptations to tool manufacture and use may be present in this species.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">It was first discovered through a clavicle bone in 2008 by nine-year-old Matthew Berger, son of paleoanthropologist Lee Berger.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Paranthropus robustus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.3 mya to 1 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kromdraai B, Swartkrans, Gondolin, Drimolen, and Coopers Cave (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">SK48 (original skull)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">410 cc to 530 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Large posterior teeth with thick enamel, consistent with other Robust Australopithecines. Enamel hypoplasia is also common in this species, possibly because of instability in the development of large, thick enameled dentition.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>P. robustus<\/em> features are neither as \u201chyper-robust\u201d as <em>P. boisei<\/em> or as ancestral in features as <em>P. aethiopicus<\/em>. They have been described as less derived, more general features that are shared with both East African species (e.g., the sagittal crest and zygomatic flaring).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Reconstructions indicate sexual dimorphism.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Several of these fossils are fragmentary in nature, distorted, and not well preserved, because they have been recovered from quarry breccia using explosives.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\"><strong><span style=\"color: #000000\">Review Questions<br \/>\n<\/span><\/strong><\/h2>\n<ul>\n<li class=\"import-Normal\"><span style=\"color: #000000\">What is the difference between a \u201cderived\u201d versus an \u201cancestral\u201d trait? Give an example of both, seen in <em>Au. afarensis<\/em>.<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">Which of the paleoenvironment hypotheses have been used to describe early hominin diversity, and which have been used to describe bipedalism?<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">Which anatomical features for bipedalism do we see in early hominins?<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">Describe the dentition of gracile and robust australopithecines. What might these tell us about their diets?<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">List the hominin species argued to be associated with stone tool technologies. Are you convinced of these associations? Why\/why not?<\/span><\/li>\n<\/ul>\n<\/div>\n<h2><span style=\"color: #000000\">Key Terms<\/span><\/h2>\n<p><span style=\"color: #000000\"><strong>Arboreal:<\/strong> Related to trees or woodland.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Aridification:<\/strong> Becoming increasingly arid or dry, as related to the climate or environment.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Aridity Hypothesis:<\/strong> The hypothesis that long-term aridification and expansion of savannah biomes were drivers in diversification in early hominin evolution.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Assemblage:<\/strong> A collection demonstrating a pattern. Often pertaining to a site or region.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Bipedalism:<\/strong> The locomotor ability to walk on two legs.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Breccia:<\/strong> Hard, calcareous sedimentary rock.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Canines:<\/strong> The pointy teeth just next to the incisors, in the front of the mouth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cheek teeth:<\/strong> Or hind dentition (molars and premolars).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Chronospecies:<\/strong> Species that are said to evolve into another species, in a linear fashion, over time.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Clade:<\/strong> A group of species or taxa with a shared common ancestor.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cladistics:<\/strong> The field of grouping organisms into those with shared ancestry.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Context:<\/strong> As pertaining to palaeoanthropology, this term refers to the place where an artifact or fossil is found.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cores:<\/strong> The remains of a rock that has been flaked or knapped.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cusps:<\/strong> The ridges or \u201cbumps\u201d on the teeth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Dental formula:<\/strong> A technique to describe the number of incisors, canines, premolars, and molars in each quadrant of the mouth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Derived traits:<\/strong> Newly evolved traits that differ from those seen in the ancestor.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Diastema:<\/strong> A tooth gap between the incisors and canines.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Early Stone Age (ESA):<\/strong> The earliest-described archaeological period in which we start seeing stone-tool technology.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>East African Rift System (EARS):<\/strong> This term is often used to refer to the Rift Valley, expanding from Malawi to Ethiopia. This active geological structure is responsible for much of the visibility of the paleoanthropological record in East Africa.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Enamel:<\/strong> The highly mineralized outer layer of the tooth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Encephalization:<\/strong> Expansion of the brain.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Extant:<\/strong> Currently living\u2014i.e., not extinct.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Fallback foods:<\/strong> Foods that may not be preferred by an animal (e.g., foods that are not nutritionally dense) but that are essential for survival in times of stress or scarcity.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Fauna:<\/strong> The animals of a particular region, habitat, or geological period.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Faunal assemblages:<\/strong> Collections of fossils of the animals found at a site.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Faunal turnover:<\/strong> The rate at which species go extinct and are replaced with new species.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Flake:<\/strong> The piece knocked off of a stone core during the manufacture of a tool, which may be used as a stone tool.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Flora:<\/strong> The plants of a particular region, habitat, or geological period.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Folivorous:<\/strong> Foliage-eating.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Foramen magnum:<\/strong> The large hole (foramen) at the base of the cranium, through which the spinal cord enters the skull.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Fossil:<\/strong> The remains or impression of an organism from the past.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Frugivorous:<\/strong> Fruit-eating.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Generalist:<\/strong> A species that can thrive in a wide variety of habitats and can have a varied diet.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Glacial:<\/strong> Colder, drier periods during an ice age when there is more ice trapped at the poles.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Gracile:<\/strong> Slender, less rugged, or pronounced features.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hallux:<\/strong> The big toe.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Holotype:<\/strong> A single specimen from which a species or taxon is described or named.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hominin:<\/strong> A primate category that includes humans and our fossil relatives since our divergence from extant great apes.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Honing P3:<\/strong> The mandibular premolar alongside the canine (in primates, the P3), which is angled to give space for (and sharpen) the upper canines.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hyper-robust:<\/strong> Even more robust than considered normal in the Paranthropus genus.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hypodigm:<\/strong> A sample (here, fossil) from which researchers extrapolate features of a population.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Incisiform:<\/strong> An adjective referring to a canine that appears more incisor-like in morphology.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Incisors:<\/strong> The teeth in the front of the mouth, used to bite off food.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Interglacial:<\/strong> A period of milder climate in between two glacial periods.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Isotopes:<\/strong> Two or more forms of the same element that contain equal numbers of protons but different numbers of neutrons, giving them the same chemical properties but different atomic masses.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Knappers:<\/strong> The people who fractured rocks in order to manufacture tools.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Knapping:<\/strong> The fracturing of rocks for the manufacture of tools.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Large Cutting Tool (LCT):<\/strong> A tool that is shaped to have functional edges.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Last Common Ancestor (LCA):<\/strong> The hypothetical final ancestor (or ancestral population) of two or more taxa before their divergence.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Lithic:<\/strong> Relating to stone (here to stone tools).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Lumbar lordosis:<\/strong> The inward curving of the lower (lumbar) parts of the spine. The lower curve in the human S-shaped spine.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Lumpers:<\/strong> Researchers who prefer to lump variable specimens into a single species or taxon and who feel high levels of variation is biologically real.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Megadont:<\/strong> An organism with extremely large dentition compared with body size.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Metacarpals:<\/strong> The long bones of the hand that connect to the phalanges (finger bones).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Molars:<\/strong> The largest, most posterior of the hind dentition.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Monophyletic:<\/strong> A taxon or group of taxa descended from a common ancestor that is not shared with another taxon or group.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Morphology:<\/strong> The study of the form or size and shape of things; in this case, skeletal parts.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Mosaic evolution:<\/strong> The concept that evolutionary change does not occur homogeneously throughout the body in organisms.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Obligate bipedalism:<\/strong> Where the primary form of locomotion for an organism is bipedal.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Occlude:<\/strong> When the teeth from the maxilla come into contact with the teeth in the mandible.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Oldowan:<\/strong> Lower Paleolithic, the earliest stone tool culture.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Orthognathic:<\/strong> The face below the eyes is relatively flat and does not jut out anteriorly.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Paleoanthropologists:<\/strong> Researchers that study human evolution.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Paleoenvironment:<\/strong> An environment from a period in the Earth\u2019s geological past.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Parabolic:<\/strong> Like a parabola (parabola-shaped).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Phalanges:<\/strong> Long bones in the hand and fingers.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Phylogenetics:<\/strong> The study of phylogeny.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Phylogeny:<\/strong> The study of the evolutionary relationships between groups of organisms.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Pliocene:<\/strong> A geological epoch between the Miocene and Pleistocene.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Polytypic:<\/strong> In reference to taxonomy, having two or more group variants capable of interacting and breeding biologically but having morphological population differences.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Postcranium:<\/strong> The skeleton below the cranium (head).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Premolars:<\/strong> The smallest of the hind teeth, behind the canines.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Procumbent:<\/strong> In reference to incisors, tilting forward.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Prognathic:<\/strong> In reference to the face, the area below the eyes juts anteriorly.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Quaternary Ice Age:<\/strong> The most recent geological time period, which includes the Pleistocene and Holocene Epochs and which is defined by the cyclicity of increasing and decreasing ice sheets at the poles.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Relative dating:<\/strong> Dating techniques that refer to a temporal sequence (i.e., older or younger than others in the reference) and do not estimate actual or absolute dates.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Robust:<\/strong> Rugged or exaggerated features.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Site:<\/strong> A place in which evidence of past societies\/species\/activities may be observed through archaeological or paleontological practice.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Specialist:<\/strong> A specialist species can thrive only in a narrow range of environmental conditions or has a limited diet.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Splitters:<\/strong> Researchers who prefer to split a highly variable taxon into multiple groups or species.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Taxa:<\/strong> Plural of taxon, a taxonomic group such as species, genus, or family.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Taxonomy:<\/strong> The science of grouping and classifying organisms.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Techno-complex:<\/strong> A term encompassing multiple assemblages that share similar traits in terms of artifact production and morphology.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Thermoregulation:<\/strong> Maintaining body temperature through physiologically cooling or warming the body.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Ungulates:<\/strong> Hoofed mammals\u2014e.g., cows and kudu.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Volcanic tufts:<\/strong> Rock made from ash from volcanic eruptions in the past.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Valgus knee:<\/strong> The angle of the knee between the femur and tibia, which allows for weight distribution to be angled closer to the point above the center of gravity (i.e., between the feet) in bipeds.<\/span><\/p>\n<h2 class=\"import-Normal\"><strong><span style=\"color: #000000\">About the Authors<br \/>\n<\/span><\/strong><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image31.jpg\" alt=\"A woman with short blonde hair smiles at the camera.\" width=\"311\" height=\"311\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Kerryn Warren, Ph.D.<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Grad Coach International, <a class=\"rId245\" style=\"color: #000000\" href=\"mailto:kerryn.warren@gmail.com\">kerryn.warren@gmail.com<\/a><\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kerryn Warren is a dissertation coach at Grad Coach International and is passionate about stimulating research thinking in students of all levels. She has lectured on multiple topics, including archaeology and human evolution, with her research and science communication interests including hybridization in the hominin fossil record (stemming from research from her Ph.D.) and understanding how evolution is taught in South African schools. She also worked as one of the \u201cUnderground Astronauts,\u201d selected to excavate <em>Homo naledi <\/em>remains from the Rising Star Cave System in the Cradle of Humankind.<\/span><\/p>\n<p>&nbsp;<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image11-2.jpg\" alt=\"A woman with short brown hair smiles at the camera.\" width=\"312\" height=\"306\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">K. Lindsay Hunter, M.A., Ph.D. candidate<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">CARTA, k.lindsay.hunter@gmail.com<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lindsay Hunter is a trained palaeoanthropologist who uses her more than 15 years of experience to make sense of the distant past of our species to build a better future. She received her master\u2019s degree in biological anthropology from the University of Iowa and is completing her Ph.D. in archaeology at the University of the Witwatersrand in Johannesburg, South Africa. She has studied fossil and human bone collections across five continents with major grant support from the National Science Foundation (United States) and the Wenner-Gren Foundation for Anthropological Research. As a National Geographic Explorer, Lindsay developed and managed the National Geographic\u2013sponsored Umsuka Public Palaeoanthropology Project in the Cradle of Humankind World Heritage Site (CoH WHS) in South Africa from within Westbury Township, Johannesburg, between 2016\u20132019. She currently serves as the Community Engagement &amp; Advancement Director for CARTA: The UC San Diego\/Salk Institute Center for Academic Research and Training in Anthropogeny in La Jolla, California.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image25-2.jpg\" alt=\"A woman with black hair stands in a hole in the ground.\" width=\"295\" height=\"339\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Navashni Naidoo, M.Sc.<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">University of Cape Town, nnaidoo2@illinois.edu<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Navashni Naidoo is a researcher at Nelson Mandela University, lecturing on physical geology. She completed her Master\u2019s in Science in Archaeology in 2017 at the University of Cape Town. Her research interests include developing paleoenvironmental proxies suited to the African continent, behavioral ecology, and engaging with community-driven archaeological projects. She has excavated at Stone Age sites across Southern Africa and East Africa. Navashni is currently pursuing a PhD in the Department of Anthropology at the University of Illinois.<\/span><\/p>\n<p>&nbsp;<\/p>\n<p>&nbsp;<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image47.jpg\" alt=\"A man with black hair and dark brown eyes looks at the camera. \" width=\"294\" height=\"294\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Silindokuhle Mavuso, M.Sc.<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">University of Witwatersrand, S.muvaso@ru.ac.za<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Silindokuhle has always been curious about the world around him and how it has been shaped. He is a lecturer at Rhodes University of Witwatersrand (Wits), and conducts research on palaeoenvironmental reconstruction and change of the northeastern Turkana Basin\u2019s Pleistocene sequence. Silindokuhle began his education with a B.Sc. (Geology, Archaeology, and Environmental and Geographical Sciences) from the University of Cape Town before moving to Wits for a B.Sc. Honors (geology and paleontology) and M.Sc. in geology. He is currently concluding his PhD Studies. During this time, he has gained more training as a Koobi Fora Fieldschool fellow (Kenya) as well as an Erasmus Mundus scholar (France). Silindokuhle is a Plio-Pleistocene geologist with a specific focus on identifying and explaining past environments that are associated with early human life and development through time. He is interested in a wide range of disciplines such as micromorphology, sedimentology, geochemistry, geochronology, and sequence stratigraphy. He has worked with teams from significant eastern and southern African hominid sites including Elandsfontein, Rising Star, Sterkfontein, Gondolin, Laetoli, Olduvai, and Koobi Fora.<\/span><\/p>\n<h2 class=\"import-Normal\"><strong><span style=\"color: #000000\">For Further Exploration<br \/>\n<\/span><\/strong><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/humanorigins.si.edu\/evidence\">The Smithsonian Institution website<\/a> hosts descriptions of fossil species, an interactive timeline, and much more.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.maropeng.co.za\/content\/page\/human-evolution\">The Maropeng Museum website<\/a> hosts a wealth of information regarding South African Fossil Bearing sites in the Cradle of Humankind<strong>.<\/strong><\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/perot-museum.imgix.net\/2019-08-naledi-sediba-quick-comparison.pdf\">This quick comparison between <em>Homo naledi<\/em> and <em>Australopithecus sediba<\/em><\/a> from the Perot Museum.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.dropbox.com\/s\/l1d2hv42psj21y9\/Braided%20Stream-1920.mp4?dl=0\">This explanation of the braided stream<\/a> by the Perot Museum.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.hetmp.com\/\">A collation of 3-D files for visualizing<\/a> (or even 3-D printing) for homes, schools, and universities.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.pbslearningmedia.org\/resource\/tdc02.sci.life.evo.lp_humanevo\/human-evolution.\">PBS learning materials<\/a>, including videos and diagrams of the Laetoli footprints, bipedalism, and fossils.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A wealth of <a href=\"https:\/\/australianmuseum.net.au\/learn\/science\/human-evolution\/\">information from the Australian Museum website<\/a>, including species descriptions, family trees, and explanations of bipedalism and diet<strong>.<\/strong><\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\"><strong>References<\/strong><\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Alemseged, Zeresenay, Fred Spoor, William H. Kimbel, Ren\u00e9 Bobe, Denis Geraads, Denn\u00e9 Reed, and Jonathan G. Wynn. 2006. \u201cA Juvenile Early Hominin Skeleton from Dikika, Ethiopia.\u201d <em>Nature<\/em> 443 (7109): 296\u2013301.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Asfaw, Berhane, Tim White, Owen Lovejoy, Bruce Latimer, Scott Simpson, and Gen Suwa. 1999. \u201c<em>Australopithecus garhi<\/em>: A New Species of Early Hominid from Ethiopia.\u201d <em>Science<\/em> 284 (5414): 629\u2013635.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Behrensmeyer, Anna K., Nancy E. Todd, Richard Potts, and Geraldine E. McBrinn. 1997. \u201cLate Pliocene Faunal Turnover in the Turkana Basin, Kenya, and Ethiopia.\u201d <em>Science<\/em> 278 (5343): 637\u2013640.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Berger, Lee R., Darryl J. De Ruiter, Steven E. Churchill, Peter Schmid, Kristian J. Carlson, Paul HGM Dirks, and Job M. Kibii. 2010. \u201c<em>Australopithecus sediba<\/em>: A New Species of <em>Homo<\/em>-like Australopith from South Africa.\u201d <em>Science<\/em> 328 (5975): 195\u2013204.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Bobe, Ren\u00e9, and Anna K. Behrensmeyer. 2004. \u201cThe Expansion of Grassland Ecosystems in Africa in Relation to Mammalian Evolution and the Origin of the Genus <em>Homo<\/em>.\u201d <em>Palaeogeography, Palaeoclimatology, Palaeoecology<\/em> 207 (3\u20134): 399\u2013420.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Brain, C. K. 1967. \u201cThe Transvaal Museum's Fossil Project at Swartkrans.\u201d <em>South African Journal of Science<\/em> 63 (9): 378\u2013384.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1938a. \u201cMore Discoveries of Australopithecus.\u201d <em>Nature<\/em> 141 (1): 828\u2013829.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1938b. \u201cThe Pleistocene Anthropoid Apes of South Africa.\u201d <em>Nature<\/em> 142 (3591): 377\u2013379.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1947. \u201cDiscovery of a New Skull of the South African Ape-Man, Plesianthropus.\u201d <em>Nature<\/em> 159 (4046): 672.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1950. \u201cThe Genera and Species of the South African Fossil Ape-Man.\u201d <em>American Journal of Physical Anthropology<\/em> 8 (1): 1\u201314.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Brunet, Michel, Alain Beauvilain, Yves Coppens, Emile Heintz, Aladji HE Moutaye, and David Pilbeam. 1995. \u201cThe First Australopithecine 2,500 Kilometers West of the Rift Valley (Chad).\u201d <em>Nature<\/em> 378 (6554): 275\u2013273.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Cerling, Thure E., Jonathan G. Wynn, Samuel A. Andanje, Michael I. Bird, David Kimutai Korir, Naomi E. Levin, William Mace, Anthony N. Macharia, Jay Quade, and Christopher H. Remien. 2011. \u201cWoody Cover and Hominin Environments in the Past 6 Million Years.\u201d <em>Nature<\/em> 476, no. 7358 (2011): 51-56..<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clarke, Ronald J. 1998. \u201cFirst Ever Discovery of a Well-Preserved Skull and Associated Skeleton of <em>Australopithecus<\/em>.\u201d <em>South African Journal of Science<\/em> 94 (10): 460\u2013463.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Clarke, Ronald J. 2013. \u201cAustralopithecus from Sterkfontein Caves, South Africa.\u201d In <em>The Paleobiology of Australopithecus<\/em>, edited by K. E. Reed, J. G. Fleagle, and R. E. Leakey, 105\u2013123. Netherlands: Springer.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clarke, Ronald J., and Kathleen Kuman. 2019. \u201cThe Skull of StW 573, a 3.67 Ma Australopithecus Prometheus Skeleton from Sterkfontein Caves, South Africa.\u201d <em>Journal of <\/em><em>H<\/em><em>uman <\/em><em>E<\/em><em>volution<\/em> 134: 102634.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clarke, R. J., and P. V. Tobias. 1995. \u201cSterkfontein Member 2 Foot Bones of the Oldest South African Hominid.\u201d <em>Science<\/em> 269 (5223): 521\u2013524.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Constantino, P. J., and B. A. Wood. 2004. \u201cParanthropus Paleobiology\u201d. In <em>Miscelanea en <\/em><em>Homenae<\/em><em> a Emiliano Aguirre<\/em><em>,<\/em> <em>v<\/em><em>olumen III: Paleoantropologia<\/em>, edited by E. G. P\u00e9rez and S. R. Jara, 136\u2013151. Alcal\u00e1 de Henares: Museo Arqueologico Regional.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Constantino, P. J., and B. A. Wood. 2007. \u201cThe Evolution of Zinjanthropus boisei.\u201d <em>Evolutionary Anthropology: <\/em><em>Issues, News, and Reviews<\/em> 16 (2): 49\u201362.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Dart, Raymond A. 1925. \u201cAustralopithecus africanus, the Man-Ape of South Africa.\u201d <em>Nature<\/em> 115: 195\u2013199.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Darwin, Charles. 1871. <em>The Descent of Man: And Selection in Relation to Sex<\/em>. London: J. Murray.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Daver, Guillaume, F. Guy, Hassane Ta\u00efsso Mackaye, Andossa Likius, J-R. Boisserie, Abderamane Moussa, Laurent Pallas, Patrick Vignaud, and N\u00e9koulnang D. Clarisse. 2022. \"Postcranial Evidence of Late Miocene Hominin Bipedalism in Chad.\" <em>Nature<\/em> 609 (7925): 94\u2013100.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Heinzelin, Jean de, J. Desmond Clark, Tim White, William Hart, Paul Renne, Giday WoldeGabriel, Yonas Beyene, and Elisabeth Vrba. 1999. \u201cEnvironment and Behavior of 2.5-Million-Year-Old Bouri Hominids.\u201d <em>Science<\/em> 284 (5414): 625\u2013629.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">DeMenocal, Peter B. D. 2004. \u201cAfrican Climate Change and Faunal Evolution during the Pliocene\u2013Pleistocene.\u201d <em>Earth and Planetary Science Letters<\/em> 220 (1\u20132): 3\u201324.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">DeMenocal, Peter B. D. and J. Bloemendal, J. 1995. \u201cPlio-Pleistocene Climatic Variability in Subtropical Africa and the Paleoenvironment of Hominid Evolution: A Combined Data-Model Approach.\u201d In <em>Paleoclimate and Evolution, with Emphasis on Human Origins<\/em>, edited by E. S. Vrba, G. H. Denton, T. C. Partridge, and L. H. Burckle, 262\u2013288. New Haven: Yale University Press.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Dirks, Paul HGM, Job M. Kibii, Brian F. Kuhn, Christine Steininger, Steven E. Churchill, Jan D. Kramers, Robyn Pickering, Daniel L. Farber, Anne-Sophie M\u00e9riaux, Andy I. R. Herries, Geoffrey C. P. King, And Lee R. Berger. 2010. \u201cGeological Setting and Age of <em>Australopithecus sediba<\/em> from Southern Africa.\u201d <em>Science<\/em> 328 (5975): 205\u2013208.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Faith, J. Tyler, and Anna K. Behrensmeyer. 2013. \u201cClimate Change and Faunal Turnover: Testing the Mechanics of the Turnover-Pulse Hypothesis with South African Fossil Data.\u201d <em>Paleobiology<\/em> 39 (4): 609\u2013627.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Grine, Frederick E. 1988. \u201cNew Craniodental Fossils of <em>Paranthropus<\/em> from the Swartkrans Formation and Their Significance in \u2018Robust\u2019 Australopithecine Evolution.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by F. E. Grine, 223\u2013243. New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Grine, Frederick E., Carrie S. Mongle, John G. Fleagle, and Ashley S. Hammond. 2022. \"The Taxonomic Attribution of African Hominin Postcrania from the Miocene through the Pleistocene: Associations and Assumptions.\" <em>Journal of Human Evolution<\/em> 173: 103255.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Haile-Selassie, Yohannes, Luis Gibert, Stephanie M. Melillo, Timothy M. Ryan, Mulugeta Alene, Alan Deino, Naomi E. Levin, Gary Scott, and Beverly Z. Saylor. 2015. \u201cNew Species from Ethiopia Further Expands Middle Pliocene Hominin Diversity.\u201d <em>Nature<\/em> 521 (7553): 432\u2013433.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Haile-Selassie, Yohannes, Stephanie M. Melillo, Antonino Vazzana, Stefano Benazzi, and Timothy M. Ryan. 2019. \u201cA 3.8-Million-Year-Old Hominin Cranium from Woranso-Mille, Ethiopia.\u201d <em>Nature<\/em> 573 (7773): 214-219.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Harmand, Sonia, Jason E. Lewis, Craig S. Feibel, Christopher J. Lepre, Sandrine Prat, Arnaud Lenoble, Xavier Bo\u00ebs et al. 2015. \u201c3.3-Million-Year-Old Stone Tools from Lomekwi3, West Turkana, Kenya.\u201d <em>Nature<\/em> 521 (7552): 310\u2013316.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Hay, Richard L. 1990. \u201cOlduvai Gorge: A Case History in the Interpretation of Hominid Paleoenvironments.\u201d In <em>East Africa: Establishment of a Geologic Framework for Paleoanthropology<\/em>, edited by L. Laporte, 23\u201337<em>.<\/em> Boulder: Geological Society of America.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Hay, Richard L., and Mary D. Leakey. 1982. \u201cThe Fossil Footprints of Laetoli.\u201d <em>Scientific American<\/em> 246 (2): 50\u201357.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Hlazo, Nomawethu. 2015. \u201cParanthropus: Variation in Cranial Morphology.\u201d Honours thesis, Archaeology Department, University of Cape Town, Cape Town.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Hlazo, Nomawethu. 2018. \u201cVariation and the Evolutionary Drivers of Diversity in the Genus <em>Paranthropus<\/em>.\u201d Master\u2019s thesis, Archaeology Department, University of Cape Town, Cape Town.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Johanson, D. C., T. D. White, and Y. Coppens. 1978. \u201cA New Species of the Genus <em>Australopithecus<\/em> (Primates: Hominidae) from the Pliocene of East Africa.\u201d <em>Kirtlandia<\/em> 28: 1\u201314.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kimbel, William H. 2015. \u201cThe Species and Diversity of Australopiths.\u201d In <em>Handbook of Paleoanthropology<\/em>, 2nd ed., edited by T. Hardt, 2071\u20132105. Berlin: Springer.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kimbel, William H., and Lucas K. Delezene. 2009. \u201c\u2018Lucy\u2019 Redux: A Review of Research on <em>Australopithecus afarensis<\/em>.\u201d <em>American <\/em><em>J<\/em><em>ournal of <\/em><em>P<\/em><em>hysical <\/em><em>A<\/em><em>nthropology<\/em> 140 (S49): 2\u201348.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Kingston, John D. 2007. \u201cShifting Adaptive Landscapes: Progress and Challenges in Reconstructing Early Hominid Environments.\u201d <em>American Journal of Physical Anthropology<\/em> 134 (S45): 20\u201358.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Kingston, John D., and Terry Harrison. 2007. \u201cIsotopic Dietary Reconstructions of Pliocene Herbivores at Laetoli: Implications for Early Hominin Paleoecology.\u201d <em>Palaeogeography, Palaeoclimatology, Palaeoecology<\/em> 243 (3\u20134): 272\u2013306.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Leakey, Louis S. B. 1959. \u201cA New Fossil Skull from Olduvai.\u201d <em>Nature<\/em> 184 (4685): 491\u2013493.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Leakey, Mary 1971. <em>Olduvai Gorge<\/em>, Vol. 3. Cambridge: Cambridge University Press.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Leakey, Mary D., and Richard L. Hay. 1979. \u201cPliocene Footprints in the Laetoli Beds at Laetoli, Northern Tanzania.\u201d <em>Nature<\/em> 278 (5702): 317\u2013323.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Leakey, Meave G., Craig S. Feibel, Ian McDougall, and Alan Walker. 1995. \u201cNew Four\u2013Million-Year-Old Hominid Species from Kanapoi and Allia Bay, Kenya.\u201d <em>Nature<\/em> 376 (6541): 565\u2013571.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Meave G., Fred Spoor, Frank H. Brown, Patrick N. Gathogo, Christopher Kiarie, Louise N. Leakey, and Ian McDougall. 2001. \u201cNew Hominin Genus from Eastern Africa Shows Diverse Middle Pliocene Lineages.\u201d <em>Nature<\/em> 410 (6827): 433\u2013440.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lebatard, Anne-Elisabeth, Didier L. Bourl\u00e8s, Philippe Duringer, Marc Jolivet, R\u00e9gis Braucher, Julien Carcaillet, Mathieu Schuster et al. 2008. \u201cCosmogenic Nuclide Dating of <em>Sahelanthropus tchadensis<\/em> and <em>Australopithecus bahrelghazali<\/em>: Mio-Pliocene Hominids from Chad.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 105 (9): 3226\u20133231.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lee-Thorp, Julia. 2011. \u201cThe Demise of \u2018Nutcracker Man.\u2019\u201d <em>Proceedings of the National Academy of Sciences<\/em> 108 (23): 9319\u20139320.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lombard, Marlize, L. Y. N. Wadley, Janette Deacon, Sarah Wurz, Isabelle Parsons, Moleboheng Mohapi, Joane Swart, and Peter Mitchell. 2012. \u201cSouth African and Lesotho Stone Age Sequence Updated.\u201d <em>The South African Archaeological Bulletin<\/em> 67 (195): 123\u2013144.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Maslin, Mark A., Chris M. Brierley, Alice M. Milner, Susanne Shultz, Martin H. Trauth, and Katy E. Wilson. 2014. \u201cEast African Climate Pulses and Early Human Evolution.\u201d <em>Quaternary Science Reviews<\/em> 101: 1\u201317.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">McHenry, Henry M. 2009. \u201cHuman Evolution.\u201d In <em>Evolution: The First Four Billion Years<\/em>, edited by M. Ruse and J. Travis, 256\u2013280. Cambridge: The Belknap Press of Harvard University Press..<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Patterson, Bryan, and William W. Howells. 1967. \u201cHominid Humeral Fragment from Early Pleistocene of Northwestern Kenya.\u201d <em>Science<\/em> 156 (3771): 64\u201366.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Pickering, Robyn, and Jan D. Kramers. 2010. \u201cRe-appraisal of the Stratigraphy and Determination of New U-Pb Dates for the Sterkfontein Hominin Site.\u201d <em>Journal of Human Evoluti<\/em><em>on<\/em> 59 (1): 70\u201386.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Potts, Richard. 1998. \u201cEnvironmental Hypotheses of Hominin Evolution.\u201d <em>American Journal of Physical Anthropology<\/em> 107 (S27): 93\u2013136.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Potts, Richard. 2013. \u201cHominin Evolution in Settings of Strong Environmental Variability.\u201d <em>Quaternary Science Reviews<\/em> 73: 1\u201313.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Rak, Yoel. 1983. <em>The <\/em><em>A<\/em><em>ustralopithecine <\/em><em>F<\/em><em>ace<\/em>. New York: Academic Press.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Rak, Yoel. 1988. \u201cOn Variation in the Masticatory System of <em>Australopithecus boisei<\/em>.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by M. Ruse and J. Travis, 193\u2013198<em>.<\/em> New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Semaw, Sileshi. 2000. \u201cThe World\u2019s Oldest Stone Artefacts from Gona, Ethiopia: Their Implications for Understanding Stone Technology and Patterns of Human Evolution between 2.6 Million Years Ago and 1.5 Million Years Ago.\u201d <em>Journal of Archaeological Science<\/em> 27(12): 1197\u20131214.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Shipman, Pat. 2002. <em>The Man Who Found the Missing Link: Eug<\/em><em>e<\/em><em>ne Dubois and <\/em><em>h<\/em><em>is Lifelong Quest to Prove Darwin Right<\/em>. New York: Simon &amp; Schuster.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Spoor, Fred. 2015. \u201cPalaeoanthropology: The Middle Pliocene Gets Crowded.\u201d<em> Nature<\/em> 521 (7553): 432\u2013433.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Strait, David S., Frederick E. Grine, and Marc A. Moniz. 1997. A Reappraisal of Early Hominid Phylogeny.\u201d <em>Journal of <\/em><em>H<\/em><em>uman <\/em><em>E<\/em><em>volution<\/em> 32 (1): 17\u201382.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Thackeray, J. Francis. 2000. \u201c\u2018Mrs. Ples\u2019 from Sterkfontein: Small Male or Large Female?\u201d <em>The South African Archaeological <\/em><em>Bulletin<\/em> 55: 155\u2013158.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Thackeray, J. Francis, Jos\u00e9 Braga, Jacques Treil, N. Niksch, and J. H. Labuschagne. 2002. \u201c\u2018Mrs. Ples\u2019 (Sts 5) from Sterkfontein: An Adolescent Male?\u201d <em>South African Journal of Science<\/em> 98 (1\u20132): 21\u201322.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Toth, Nicholas. 1985. \u201cThe Oldowan Reassessed.\u201d <em>Journal of Archaeological Science<\/em>\u00a012 (2): 101\u2013120.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Vrba, E. S. 1988. \u201cLate Pliocene Climatic Events and Hominid Evolution.\u201d In <em>The <\/em><em>E<\/em><em>volutionary <\/em><em>H<\/em><em>istory of the <\/em><em>R<\/em><em>obust Australopithecines<\/em>, edited by F. E. Grine, 405\u2013426. New York: Aldine.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Vrba, Elisabeth S. 1998. \u201cMultiphasic Growth Models and the Evolution of Prolonged Growth Exemplified by Human Brain Evolution.\u201d <em>Journal of Theoretical Biology<\/em> 190 (3): 227\u2013239.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Vrba, Elisabeth S. 2000. \u201cMajor Features of Neogene Mammalian Evolution in Africa.\u201d In <em>Cenozoic <\/em><em>G<\/em><em>eology of <\/em><em>S<\/em><em>outhern Africa<\/em>, edited by T. C. Partridge and R. Maud, 277\u2013304<em>.<\/em> Oxford: Oxford University Press.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Walker, Alan C., and Richard E. Leakey. 1988. \u201cThe Evolution of <em>Australopithecus boisei<\/em>.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by F. E. Grine, 247\u2013258. New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Walker, Alan, Richard E. Leakey, John M. Harris, and Francis H. Brown. 1986. \u201c2.5-my <em>Australopithecus boisei<\/em> from West of Lake Turkana, Kenya.\u201d <em>Nature<\/em> 322 (6079): 517\u2013522.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Ward, Carol, Meave Leakey, and Alan Walker. 1999. \u201cThe New Hominid Species <em>Australopithecus anamensis<\/em>.\u201d <em>Evolutionary Anthropology<\/em> 7 (6): 197\u2013205.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">White, Tim D. 1988. \u201cThe Comparative Biology of \u2018Robust\u2019 Australopithecus: Clues from Content.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by F. E. Grine, 449\u2013483. New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">White, Tim D., Gen Suwa, and Berhane Asfaw. 1994. \u201c<em>Australopithecus ramidus<\/em>, a New Species of Early Hominid from Aramis, Ethiopia.\u201d <em>Nature<\/em> 371 (6495): 306\u2013312.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Wood, Bernard. 2010. \u201cReconstructing Human Evolution: Achievements, Challenges, and Opportunities.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 10 (2): 8902\u20138909.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Wood, Bernard, and Eve K. Boyle. 2016. \u201cHominin Taxic Diversity: Fact or Fantasy?\u201d <em>Yearbook of Physical Anthropology<\/em> 159 (S61): 37\u201378.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Wood, Bernard, and Kes Schroer. 2017. \u201cParanthropus: Where Do Things Stand?\u201d In <em>Human Paleontology and Prehistory<\/em>, edited by A. Marom and E. Hovers, 95\u2013107. New York: Springer, Cham.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Acknowledgements<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">All of the authors in this section are students and early career researchers in paleoanthropology and related fields in South Africa (or at least have worked in South Africa). We wish to thank everyone who supports young and diverse talent in this field and would love to further acknowledge Black, African, and female academics who have helped pave the way for us.<\/span><\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1770\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1770\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Sarah S. King, Ph.D., Cerro Coso Community College<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Kara Jones, M.A., Ph.D. student, University of Nevada Las Vegas<\/p>\n<p class=\"import-Normal\"><em>This chapter<\/em><em> is a revision from \"<\/em><a class=\"rId6\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-7\/\"><em>Chapter 7: Understanding the Fossil Context<\/em><\/a><em>\u201d by Sarah King and Lee Anne Zajicek. <\/em><em>In <\/em><a class=\"rId7\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, first edition<\/em><\/a><em>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under <\/em><a class=\"rId8\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\"><em>CC BY-NC 4.0<\/em><\/a><em>. <\/em><\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\"><span style=\"color: #ffffff;\">Learning Objectives<\/span><\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li class=\"import-Normal\" style=\"text-indent: 0pt;\">Identify the different types of fossils and describe how they are formed.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt;\">Discuss relative and chronometric dating methods, the type of material they analyze, and their applications.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt;\">Describe the methods used to reconstruct past environments.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt;\">Interpret a site using the methods described in this chapter.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Fossil Study: An Evolving Process<\/h2>\n<h3 class=\"import-Normal\"><strong>Mary Anning and the Age of Wonder<\/strong><\/h3>\n<figure style=\"width: 206px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/05\/image12.jpg\" alt=\"Woman points to dog and fossil on the ground.\" width=\"206\" height=\"248\" \/><figcaption class=\"wp-caption-text\">Figure 7.1: An oil painting of Mary Anning and her dog, Tray, prior to 1845. The \u201cJurassic Coast\u201d of Lyme Regis is in the background. Notice that Anning is pointing at a fossil. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Mary_Anning_by_B._J._Donne.jpg\">Mary Anning by B. J. Donne<\/a> from the Geological Society\/NHMPL is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p>Mary Anning (1799\u20131847) is likely the most famous fossil hunter you\u2019ve never heard of (Figure 7.1). Anning lived her entire life in Lyme Regis on the Dorset coast in England. As a woman, born to a poor family, with minimal education (even by 19th-century standards), the odds were against Anning becoming a scientist (Emling 2009, xii). It was remarkable that Anning was eventually able to influence the great scientists of the day with her fossil discoveries and her subsequent hypotheses regarding evolution.<\/p>\n<p class=\"import-Normal\">The time when Anning lived was a remarkable period in human history because of the Industrial Revolution in Britain. Moreover, the scientific discoveries of the 18th and 19th centuries set the stage for great leaps of knowledge and understanding about humans and the natural world. Barely a century earlier, Sir Isaac Newton had developed his theories on physics and become the president of the Royal Society of London (Dolnick 2011, 5). In this framework, the pursuit of intellectual and scientific discovery became a popular avocation for many individuals, the vast majority of whom were wealthy men (Figure 7.2).<\/p>\n<figure style=\"width: 358px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image22-1.png\" alt=\"Robed figure near a rock structure.\" width=\"358\" height=\"273\" \/><figcaption class=\"wp-caption-text\">Figure 7.2: A Walk at Dusk, 1830\u20131835, by Caspar David Friedrich, is a painting likely of a dolmen, a megalithic (large rock) tomb. Dolmens were built throughout Europe, five to six thousand years ago. Scholars were fascinated by the ancient world, which was an accepted part of Earth\u2019s history, even if explanation defied nonsecular thought. Credit: <a href=\"https:\/\/www.getty.edu\/art\/collection\/object\/103RJX\">A Walk at Dusk object 93.PA.14<\/a> by Casper David Friedrich German, 1774\u20131840, Paul Getty Museum, is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a> and part of the <a href=\"https:\/\/www.getty.edu\/projects\/open-content-program\/\">Getty Open Content Program<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">In spite of the expectations of Georgian English society to the contrary, Anning became a highly successful fossil hunter as well as a self-educated geologist and anatomist. The geology of Lyme Regis, with its limestone cliffs, provided a fortuitous backdrop for Anning\u2019s lifework. Now called the \u201cJurassic Coast,\u201d Lyme Regis has always been a rich source for fossilized remains (Figure 7.3). Continuing her father\u2019s passion for fossil hunting, Anning scoured the crumbling cliffs after storms for fossilized remains and shells. The work was physically demanding and downright dangerous. In 1833, while searching for fossils, Anning lost her beloved dog in a landslide and nearly lost her own life in the process (Emling 2009).<\/p>\n<figure style=\"width: 283px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image14-1.jpg\" alt=\"Rocky coastline and cliffs.\" width=\"283\" height=\"212\" \/><figcaption class=\"wp-caption-text\">Figure 7.3: The \u201cJurassic Coast\u201d of Lyme Regis: the home of fossil hunter Mary Anning. Credit: <a href=\"https:\/\/pixabay.com\/photos\/lyme-regis-coast-sea-cliffs-924431\/\">Lyme-regis-coast-sea-cliffs-924431<\/a> by <a href=\"https:\/\/pixabay.com\/users\/jstarj-884623\/\">jstarj<\/a> has been designated to the <a href=\"https:\/\/creativecommons.org\/share-your-work\/public-domain\/cc0\/\">public domain (CC0)<\/a> under a <a href=\"https:\/\/pixabay.com\/service\/terms\/#license\">Pixabay License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Around the age of ten, Anning located and excavated a complete fossilized skeleton of an ichthyosaurus (\u201cfish lizard\u201d). She eventually found <em>Pterodactylus macronyx<\/em> and a 2.7-meter <em>Plesiosaurus<\/em>, considered by many to be her greatest discovery (Figure 7.4). These discoveries proved that there had been significant changes in the way living things appeared throughout the history of the world. Like many of her peers, including Darwin, Anning had strong religious convictions. However, the evidence that was being found in the fossil record was contradictory to the Genesis story in the Bible. In <em>The Fossil Hunter: Dinosaurs, Evolution, and the Woman Whose Discoveries Changed the World<\/em>, Anning\u2019s biographer Shelley Emling (2009, 38) notes, \u201cthe puzzling attributes of Mary\u2019s fossil [ichthyosaurus] struck a blow at this belief and eventually helped pave the way for a real understanding of life before the age of humans.\u201d<\/p>\n<figure style=\"width: 247px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image21.png\" alt=\"Plesiosaurus drawing.\" width=\"247\" height=\"375\" \/><figcaption class=\"wp-caption-text\">Figure 7.4: Plesiosaurus, illustrated and described by Mary Anning in an undated handwritten letter. Credit: <a href=\"https:\/\/wellcomecollection.org\/works\/cezbevj4\">Autograph letter concerning the discovery of plesiosaurus<\/a> by Mary Anning (1799\u20131847) from the <a href=\"https:\/\/wellcomecollection.org\">Wellcome Collection<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/\">CC BY 4.0 License<\/a>.<\/figcaption><\/figure>\n<p>Intellectual and scientific debate now had physical evidence to support the theory of evolution, which would eventually result in Darwin\u2019s seminal work,<em> On the Origin of Species<\/em> (1859). Anning\u2019s discoveries and theories were appreciated and advocated by her friends, intellectual men who were associated with the Geological Society of London. Regrettably, this organization was closed to women, and Anning received little official recognition for her contributions to the fields of natural history and paleontology. It is clear that Anning\u2019s knowledge, diligence, and uncanny luck in finding magnificent specimens of fossils earned her unshakeable credibility and made her a peer to many antiquarians (Emling 2009).<\/p>\n<p class=\"import-Normal\">Fossil hunting is still providing evidence and a narrative of the story of Earth. Mary Anning recognized the value of fossils in understanding natural history and relentlessly championed her theories to the brightest minds of her day. Anning\u2019s ability to creatively think \u201coutside the box\u201d\u2014skillfully assimilating knowledge from multiple academic fields\u2014was her gift to our present understanding of the fossil record. Given how profoundly Anning has shaped how we, in the modern day, think about the origins of life, it is surprising that her contributions have been so marginalized. Anning\u2019s name should be on the tip of everyone\u2019s tongue. Fortunately, at least in one sense of the word, it is. The well-known tongue twister, below, may have been written about Mary Anning:<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 130.5pt; text-indent: 36pt;\">She sells sea-shells on the sea-shore.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 130.5pt; text-indent: 36pt;\">The shells she sells are sea-shells, I\u2019m sure.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 130.5pt; text-indent: 36pt;\">For if she sells sea-shells on the sea-shore<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 130.5pt; text-indent: 36pt;\">Then I\u2019m sure she sells sea-shore shells.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 130.5pt; text-indent: 36pt;\">\u2014T. Sullivan (1908)<\/p>\n<h3 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Developing Modern <\/strong><strong>Methods<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">As Mary Anning\u2019s story suggests, scientists in Europe were working at a time dominated by western Christian tradition. Literal interpretations of the bible did not allow for the long, slow processes of geological or evolutionary change to operate. However, many scientists were making observations that did not fit the biblical narrative. During the 18th century, Scotsman James Hutton\u2019s work on the formation of Earth provided a much longer timeline of events than previous biblical interpretations would allow. Hutton\u2019s theory of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_826\">Deep Time<\/a><\/strong> was crucial to the understanding of fossils. Deep Time gave the history of Earth enough time\u20144.543 billion years\u2014to encompass <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_828\">continental drift<\/a><\/strong>, the evolution of species, and the fossilization process. A second Scotsman, Charles Lyell, propelled Hutton\u2019s work into his own theory of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_830\">uniformitarianism<\/a><\/strong>, the doctrine that Earth\u2019s geologic formations are the work of slow geologic forces. Lyell\u2019s three-volume work, <em>Principles of Geology<\/em> (1830\u20131833), was influential to naturalist Charles Darwin (see Chapter 2 for more information on Darwin\u2019s work). In fact, Lyell\u2019s first volume accompanied Darwin on his five-year voyage around the world on the <em>HMS Beagle<\/em> (1831\u20131836). The concepts proposed by Lyell gave Darwin an opportunity to apply his working theories of evolution by natural selection and a greater length of time with which to work. These resulting theories were important scientific discoveries and paved the way for the \u201cAge of Wonder\u201d (Holmes 2010, xvi).<\/p>\n<figure style=\"width: 264px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image30-1.jpg\" alt=\"Fossilized shell.\" width=\"264\" height=\"176\" \/><figcaption class=\"wp-caption-text\">Figure 7.5: Murexsul (Miocene): This fossil was found at the Naval Weapons Center, China Lake, California, in 1945. The fossil was buried deep in the strata and was pulled out of the ground along with a crashed \u201cFat Boy\u201d missile after atomic missile testing (S. Brubaker, personal communication, March 9, 2018). Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-7\/\">Murexsul (Figure 7.6)<\/a> from the <a href=\"https:\/\/maturango.org\/\">Maturango Museum<\/a>, Ridgecrest, California, by Sarah S. King and Lee Anne Zajicek is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p>The work of Anning, Darwin, Lyell, and many others laid the foundation for the modern methods we use today. Though anthropology is focused on humans and our primate relatives (and not on dinosaurs, as many people wrongly assume), you will see that methods developed in paleontology, geology, chemistry, biology, and physics are often applied in anthropological research. In this chapter, you will learn about the primary methods and techniques employed by biological anthropologists to answer questions about <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_832\">fossils<\/a><\/strong>, the mineralized copies of once-living organisms (Figure 7.5). Ultimately, these answers provide insights into human evolution. Pay close attention to ways in which modern biological anthropologists use other disciplines to analyze evidence and reconstruct past activities and environments.<\/p>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Earth: It's Older than Dirt<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Scientists have developed precise and accurate dating methods based on work in the fields of physics and chemistry. Using these methods, scientists are able to establish the age of Earth as well as approximate ages of the organisms that have lived here. Earth is roughly 4.6 billion years old, give or take a few hundred million years. The first evidence for a living organism appeared around 3.5 billion years ago (<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_844\">bya<\/a><\/strong>)<strong>.<\/strong> The scale of geologic time can seem downright overwhelming. In order to organize and make sense of Earth\u2019s past, geologists break up that time into subunits, which are human-made divisions along Earth\u2019s timeline. The largest subunit is the <strong>eon. <\/strong>An eon is further divided into <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_836\">eras<\/a>,<\/strong> and eras are divided into <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_838\">periods<\/a><\/strong>. Finally, periods are divided into <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_846\">epochs<\/a><\/strong> (see Figure 7.6; Williams 2004, 37). Currently, we are living in the Phanerozoic eon, Cenozoic era, Quaternary period, and probably the Holocene epoch\u2014though there is academic debate about the current epoch (see below).<\/p>\n<figure id=\"attachment_248\" aria-describedby=\"caption-attachment-248\" style=\"width: 1134px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-227 size-full\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/Geo-Time-Scale-FY17.jpeg\" alt=\"Table of geological time scale and examples. Full text link in caption.\" width=\"1134\" height=\"1300\" \/><figcaption id=\"caption-attachment-248\" class=\"wp-caption-text\">Figure 7.6: The Geologic time scale is shown here, with periods broken into eons, eras, periods, and in some cases epochs. Some life forms and geological events are noted for each period. <a href=\"https:\/\/docs.google.com\/document\/d\/1VUDKMBJYS_jNONjLxT04jQN0_z9Ua50BRN6auGSHUuU\/edit\">A full text description of this image is available<\/a>. Credit: <a href=\"https:\/\/www.nps.gov\/subjects\/geology\/time-scale.htm\" target=\"_blank\" rel=\"noopener\">Geologic Time Scale<\/a>, by <a href=\"https:\/\/www.nps.gov\/index.htm\" target=\"_blank\" rel=\"noopener\">National Park Service<\/a>, designed by Trista Thornberry-Ehrlich and Rebecca Port, adapted from ones from <a href=\"https:\/\/www.usgs.gov\/\" target=\"_blank\" rel=\"noopener\">USGS<\/a> and the International Commission on Stratigraphy, is in the <a href=\"https:\/\/www.nps.gov\/aboutus\/disclaimer.htm#:~:text=%C2%A7%C2%A7%20101%2C%20105)\" target=\"_blank\" rel=\"noopener\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">These divisions are based on major changes and events recorded in the geologic record. Events like significant shifts in climate or mass extinctions can be used to mark the end of one geologic time unit and the beginning of another. However, it is important to remember that these borders are not real in a physical sense; they are helpful organizational guidelines for scientific research. There can be debate regarding how the boundaries are defined. Additionally, the methods we use to establish these dates are refined over time, occasionally leading to shifts in established chronology (see the discussion on calibration in the radiocarbon dating section below). For instance, the current epoch has been traditionally known as the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_840\">Holocene<\/a><\/strong>. It began almost twelve thousand years ago (<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_842\">kya<\/a><\/strong>) during the warming period after that last major ice age. Today, there is evidence to indicate human-driven climate change is warming the world and changing the environmental patterns faster than the natural cyclical processes. This has led some scientists within the stratigraphic community to argue for a new epoch beginning around 1950 with the Nuclear Age called the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_848\">Anthropocene<\/a> <\/strong>(Monastersky 2015; Waters et al. 2016). Nobel Laureate Paul Crutzen places the beginning of the Anthropocene much earlier\u2014at the dawn of the Industrial Revolution, with its polluting effects of burning coal (Crutzen and Stoermer 2000, 17\u201318). Geologist William Ruddiman argues that the epoch began 5,000\u20138,000 years ago with the advent of agriculture and the buildup of early methane gasses (Ruddiman et al. 2008). Regardless of when the Anthropocene started, the major event that marks the boundary is the warming temperatures and mass extinction of nonhuman species caused by human activity (Figure 7.7). Researchers now declare that \u201chuman activity now rivals geologic forces in influencing the trajectory of the Earth System\u201d (Steffen et al. 2018, 1).<\/p>\n<figure style=\"width: 299px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image1.jpg\" alt=\"Two cylindrical towers emitting white steam.\" width=\"299\" height=\"168\" \/><figcaption class=\"wp-caption-text\">Figure 7.7: The Chooz Nuclear Power, in a valley in Ardennes, France, is a reminder that human activity affects the planet greatly. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Chooz_Nuclear_Power_Plant-9361.jpg\">Chooz Nuclear Power Plant-9361<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Raymond\">Raimond Spekking<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Fossils: The Taphonomic Process<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Most of the evidence of human evolution comes from the study of the dead. To obtain as much information as possible from the remains of once-living creatures, one must understand the processes that occur after death. This is where <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_850\">taphonomy<\/a><\/strong> comes in (Figure 7.8). Taphonomy is the study of what happens to an organism after death (Komar and Buikstra 2008, 189; Stodder 2008). It includes the study of how an organism becomes a fossil. However, as you\u2019ll see throughout this book, the majority of organisms never make it through the full fossilization process.<\/p>\n<figure style=\"width: 261px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image25-1.jpg\" alt=\"Coyote skull with bones and fur.\" width=\"261\" height=\"348\" \/><figcaption class=\"wp-caption-text\">Figure 7.8: Taphonomy focuses on what happens to the remains of an organism, like this coyote, after death. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-7\/\">Coyote remains (Figure 7.14)<\/a> by Sarah S. King is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Taphonomy is important in biological anthropology, especially in subdisciplines like bioarchaeology (the study of human remains in the archaeological record) and zooarchaeology (the study of faunal remains from archaeological sites). It is so important that many scientists have recreated a variety of burial and decay experiments to track taphonomic change in modern contexts. These contexts can then be used to understand the taphonomic patterns seen in the fossil record (see Reitz and Wing 1999, 122\u2013141).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Taphonomic analysis can also give us important insights into the development of complex thought and ritual in human evolution. In Chapter 11, you will see the first evidence of recognized burial practices in hominins. Taphonomy helped to establish whether these burials were simply the result of natural processes or intentionally constructed by humans (Klein 1999, 395; Straus 1989). Deliberate burials often include the body placed in a specific position, such as supine (on the back) with arms crossed over the chest or in a flexed position (think fetal position) facing a particular direction. If bones have evidence of a carnivore or rodent gnawing on them, it can be inferred that the remains were exposed to scavengers after death.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Going back further in time, taphonomic evidence may tell us how our ancestors died. For instance, several australopithecine fossils show evidence of carnivore tooth marks and even punctures from saber-toothed cats, indicating that we weren\u2019t always the top of the food chain. The Bodo Cranium, a <em>Homo erectus<\/em> cranium from Middle Awash Valley, Ethiopia, shows cut marks made by stone tools, indicating an early example of possible defleshing activity in our human ancestors (White 1986). At the archaeological site of Zhoukoudian, researchers used taphonomy to show that the highly fragmented remains of at least 51 <em>Homo erectus<\/em> individuals were scavenged by Pleistocene cave hyenas (Boaz et al. 2004). The damage on Skull VI was described as \u201celongated, raking bite marks, isolated puncture bite marks, and perimortem breakage consistent with patterns of modern hyaenid bone modification\u201d (Boaz et al. 2004). Additionally, a fresh burnt equid cranium was discovered which supports the theory of mobile hominid scavenging and fire use at the site (Boaz et al. 2004).<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Special Topic: Bog Bodies and Mummies <span style=\"text-decoration: underline;\">(Add a picture to show how well preserved they are!)<\/span><\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Preservation is a key topic in anthropological research, since we can only study the evidence that gets left behind in the fossil and archaeological record. This chapter is concerned with the fossil record; however, there are other forms of preserved remains that provide anthropologists with information about the past. You\u2019ve undoubtedly heard of mummification, likely in the context of Egyptian or South American mummies. However, bog bodies and ice mummies are further examples of how remains can be preserved in special circumstances. It is important to note that fossilization is a process that takes much longer than the preservation of bog bodies or mummies.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Bog bodies are good examples of wetland preservation. Peat bogs are formed by the slow accumulation of vegetation and silts in ponds and lakes. Individuals were buried in bogs throughout Europe as far back as 10 kya, with a proliferation of activity from 1,600 to 3,200 years ago (Giles 2020; Ravn 2010). When they were found thousands of years later, they resembled recent burials. Their hair, skin, clothing, and organs were exceptionally well preserved, in addition to their bones and teeth (Eisenbeiss 2016; Ravn 2010). Preservation was so good in fact that archaeologists could identify the individuals\u2019 last meals and re-create tattoos found on their skin.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Extreme cold can also halt the natural decay process. A well-known ice mummy is \u00d6tzi, a Copper Age man dating to around 5,200 years ago found in the Alps (Vanzetti et al. 2012; Vidale et al. 2016). As with the bog bodies, his hair, skin, clothing, and organs were all well preserved. Recently, archaeologists were able to identify his last meal (Maixner et al. 2018). It was high in fat, which makes sense considering the extremely cold environment in which he lived, as meals high in fat assist in cold tolerance (Fumagalli et al. 2015).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">In the Andes, ancient peoples would bury human sacrifices throughout the high peaks in a sacred ritual called Capacocha (Wilson et al. 2007). The best-preserved mummy to date is called the \u201cMaiden\u201d or \u201cSarita\u201d because she was found at the summit of Sara Sara Volcano. Her remains are over 500 years old, but she still looks like the 15-year-old girl she was at the time of her death, as if she had just been sleeping for 500 years (Reinhard 2006).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Finally, arid environments can also contribute to the preservation of organic remains. As discussed with waterlogged sites, much of the bacteria that is active in breaking down bodies is already present in our gut and begins the putrefaction process shortly after death. Arid environments deplete organic material of the moisture that putrefactive bacteria need to function (Booth et al. 2015). When that occurs, the soft tissue like skin, hair, and organs can be preserved. It is similar to the way a food dehydrator works to preserve meat, fruit, and vegetables for long-term storage. There are several examples of arid environments spontaneously preserving human remains, including catacomb burials in Austria and Italy (Aufderheide 2003, 170, 192\u2013205).<\/p>\n<\/div>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Fossilization<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Fossils only represent a tiny fraction of creatures that existed in the past. It is extremely difficult for an organism to become a fossil. After all, organisms are designed to deteriorate after they die. Bacteria, insects, scavengers, weather, and environment all aid in the process that breaks down organisms so their elements can be returned to Earth to maintain ecosystems (Stodder 2008). <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_852\">Fossilization<\/a><\/strong>, therefore, is the preservation of an organism against these natural decay processes (Figure 7.9).<\/p>\n<figure style=\"width: 699px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image20-2.png\" alt=\"Five images depicting fossilization.\" width=\"699\" height=\"345\" \/><figcaption class=\"wp-caption-text\">Figure 7.9: A simplified illustration of the fossilization process beginning at an organism's death. In this example, the individual begins to decompose and then is covered by water and sediments, both protecting it and creating an environment for perimineralization. Sediments accumulate over time. Erosion eventually exposes the fossil, leading to its eventual discovery by paleoanthropologists. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-7\/\">Fossilization process (Figure 7.15)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">For fossilization to occur, several important things must happen. First, the organism must be protected from things like bacterial activity, scavengers, and temperature and moisture fluctuations. A stable environment is important. This means that the organism should not be exposed to significant fluctuations in temperature, humidity, and weather patterns. Changes to moisture and temperature cause the organic tissues to expand and contract repeatedly, which will eventually cause microfractures and break down (Stodder 2008). Soft tissue like organs, muscle, and skin are more easily broken down in the decay process; therefore, they are less likely to be preserved. Bones and teeth, however, last much longer and are more common in the fossil record (Williams 2004, 207).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Wetlands are a particularly good area for preservation because they allow for rapid permanent burial and a stable moisture environment. That is why many fossils are found in and around ancient lakes and river systems. Waterlogged sites can also be naturally <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_854\">anaerobic<\/a><\/strong> (without oxygen). Much of the bacteria that causes decay is already present in our gut and can begin the decomposition process shortly after death during putrefaction (Booth et al. 2015). Since oxygen is necessary for the body\u2019s bacteria to break down organic material, the decay process is significantly slowed or halted in anaerobic conditions.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">The next step in the fossilization process is sediment accumulation. The sediments cover and protect the organism from the environment. They, along with water, provide the minerals that will eventually become the fossil (Williams 2004, 31). Sediment accumulation also provides the pressure needed for mineralization to take place. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_856\">Lithification<\/a><\/strong> is when the weight and pressure of the sediments squeeze out extra fluids and replace the voids that appear with minerals from the surrounding sediments. Finally, we have <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_858\">permineralization<\/a><\/strong>. This is when the organism is fully replaced by minerals from the sediments. A fossil is really a mineral copy of the original organism (Williams 2004, 31).<\/p>\n<h3 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Types of Fossils<\/strong><\/h3>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><em>Plants<\/em><\/h4>\n<figure style=\"width: 259px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image2-1.jpg\" alt=\"Petrified wood.\" width=\"259\" height=\"194\" \/><figcaption class=\"wp-caption-text\">Figure 7.10: An exquisite piece of petrified wood. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:PetrifiedWood.jpg\">PetrifiedWood<\/a> at the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Petrified_Forest_National_Park\">Petrified Forest National Park<\/a> by <a href=\"https:\/\/pdphoto.org\/\">Jon Sullivan<\/a> has been designated to the <a href=\"https:\/\/creativecommons.org\/share-your-work\/public-domain\/cc0\/\">public domain (CC0)<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Plants make up the majority of fossilized materials. One of the most common plants existing today, the fern, has been found in fossilized form many times. Other plants that no longer exist or the early ancestors of modern plants come in fossilized forms as well. It is through these fossils that we can discover how plants evolved and learn about the climate of Earth over different periods of time.<\/p>\n<p>Another type of fossilized plant is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_860\">petrified wood<\/a><\/strong>. This fossil is created when actual pieces of wood\u2014such as the trunk of a tree\u2014mineralize and turn into rock. Petrified wood is a combination of silica, calcite, and quartz, and it is both heavy and brittle. Petrified wood can be colorful and is generally aesthetically pleasing because all the features of the original tree\u2019s composition are illuminated through mineralization (Figure 7.10). There are a number of places all over the world where petrified wood \u201cforests\u201d can be found, but there is an excellent assemblage in Arizona, at the Petrified Forest National Park. At this site, evidence relating to the environment of the area some 225 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_862\">mya<\/a><\/strong> is on display.<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><em>Human\/Animal Remains<\/em><\/h4>\n<figure style=\"width: 242px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image7-1.jpg\" alt=\"Partial hominin skeleton on black background.\" width=\"242\" height=\"583\" \/><figcaption class=\"wp-caption-text\">Figure 7.11: \u201cLucy\u201d (AL 288-1), Australopithecus afarensis. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Lucy_blackbg.jpg\">Lucy blackbg<\/a> by 120 is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.5\/deed.en\">CC BY 2.5 License<\/a>.<\/figcaption><\/figure>\n<p>We are more familiar with the fossils of early animals because natural history museums have exhibits of dinosaurs and extinct mammals. However, there are a number of fossilized hominin remains that provide a picture of the fossil record over the course of our evolution from primates. The term <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_800\">hominins<\/a><\/strong> includes all human ancestors who existed after the evolutionary split from chimpanzees and bonobos, some six to seven mya. Modern humans are <em>Homo sapiens<\/em>, but hominins can include much earlier versions of humans. One such hominin is \u201cLucy\u201d (AL 288-1), the 3.2 million-year-old fossil of <em>Australopithecus afarensis<\/em> that was discovered in Ethiopia in 1974 (Figure 7.11). Until recently, Lucy was the most complete and oldest hominin fossil, with 40% of her skeleton preserved (see Chapter 9 for more information about Lucy). In 1994, an <em>Australopithecus<\/em> fossil nicknamed \u201cLittle Foot\u201d (Stw 573) was located in the World Heritage Site at Sterkfontein Caves (\u201cthe Cradle of Humankind\u201d) in South Africa. Little Foot is more complete than Lucy and possibly the oldest fossil that has so far been found, dating to at least 3.6 million years (Granger et al. 2015). The ankle bones of the fossil were extricated from the matrix of concrete-like rock, revealing that the bones of the ankles and feet indicate bipedalism (University of Witwatersrand 2017).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Both the Lucy and Little Foot fossils date back to the Pliocene (5.8 to 2.3 mya). Older hominin fossils from the late Miocene (7.25 to 5.5 mya) have been located, although they are much less complete. The oldest hominin fossil is a fragmentary skull named <em>Sahelanthropus tchadensis<\/em>, found in Northern Chad and dating to circa seven mya (Lebatard et al. 2008). It is through the discovery, dating, and study of primate and early hominin fossils that we find physical evidence of the evolutionary timeline of humans.<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><em>Asphalt<\/em><\/h4>\n<figure style=\"width: 510px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image28.jpg\" alt=\"Asphalt lake with mammoth figurines.\" width=\"510\" height=\"340\" \/><figcaption class=\"wp-caption-text\">Figure 7.12: This is a recreation of how animals tragically came to be trapped in the asphalt lake at the La Brea Tar Pits. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Mammoth_Tragedy_at_La_Brea_Tar_Pits_(5463657162).jpg\">Mammoth Tragedy at La Brea Tar Pits (5463657162)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/81943113@N00\">KimonBerlin<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/legalcode\">CC BY-SA 2.0 License<\/a>.<\/figcaption><\/figure>\n<figure style=\"width: 206px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image6-3.jpg\" alt=\"Skull with open jaw and large teeth.\" width=\"206\" height=\"245\" \/><figcaption class=\"wp-caption-text\">Figure 7.13: The fearsome jaws of the saber-toothed cat (Smilodon fatalis) found at the La Brea Tar Pits. Credit: <a href=\"https:\/\/www.flickr.com\/photos\/jsjgeology\/15256884929\">Smilodon saber-toothed tiger skull (La Brea Asphalt, Upper Pleistocene; Rancho La Brea tar pits, southern California, USA) 1<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/jsjgeology\/\">James St. John<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\">CC BY 2.0 License<\/a>.<\/figcaption><\/figure>\n<p>Asphalt, a form of crude oil, can also yield fossilized remains. Asphalt is commonly referred to in error as tar because of its viscous nature and dark color. A famous fossil site from California is La Brea Tar Pits in downtown Los Angeles (Figure 7.12). In the middle of the busy city on Wilshire Boulevard, asphalt (not tar) bubbles up through seeps (cracks) in the sidewalk. The La Brea Tar Pits Museum provides an incredible look at the both extinct and extant animals that lived in the Los Angeles Basin 40,000\u201311,000 years ago. These animals became entrapped in the asphalt during the Pleistocene and perished in place. Ongoing excavations have yielded millions of fossils, including <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_864\">megafauna<\/a><\/strong> such as American mastodons and incomplete skeletons of extinct species of dire wolves, <em>Canis dirus<\/em>, and the saber-toothed cat, <em>Smilodon fatalis<\/em> (Figure 7.13). Fossilized remains of plants have also been found in the asphalt. The remains of one person have also been found at the tar pits. Referred to as La Brea Woman, the remains were found in 1914 and were subsequently dated to around 10,250 years ago. The La Brea Woman was a likely female individual who was 17\u201328 years old at the time of her death, with a height of under five feet (Spray 2022). She is thought to have died from blunt force trauma to her head, famously making her Los Angeles\u2019s first documented homicide victim (Spray 2022). (Learn more about her in the Special Topic box, \u201cNecropolitics,\u201d below.) Between the fossils of animals and those of plants, paleontologists have a good idea of the way the Los Angeles Basin looked and what the climate in the area was like many thousands of years ago.<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><em>Igneous Rock<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Most fossils are found in sedimentary rock. This type of rock has been formed from deposits of minerals over millions of years in bodies of water on Earth\u2019s surface. Some examples include shale, limestone, and siltstone. Sedimentary rock typically has a layered appearance. However, fossils have been found in igneous rock as well. Igneous rock is volcanic rock that is created from cooled molten lava. It is rare for fossils to survive molten lava, and it is estimated that only 2% of all fossils have been found in igneous rock (Ingber 2012). Part of a giant rhinocerotid skull dating back 9.2 mya to the Miocene was discovered in Cappadocia, Turkey, in 2010. The fossil was a remarkable find because the eruption of the \u00c7ardak caldera was so sudden that it simply dehydrated and \u201cbaked\u201d the animal (Antoine et al. 2012).<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><em>Trace Fossils<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Depending on the specific circumstances of weather and time, even footprints can become fossilized. Footprints fall into the category of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_866\">trace fossils<\/a><\/strong>, which includes other evidence of biological activity such as nests, burrows, tooth marks, and shells. A well-known example of trace fossils are the Laetoli footprints in Tanzania (Figure 7.14). More recently, archaeological investigations in North America have revealed fossil footprints which rewrite the history of people in the Americas at White Sands, New Mexico. You can read more about the Laetoli and White Sands footprints in the Dig Deeper box below.<\/p>\n<figure style=\"width: 399px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image5-2.jpg\" alt=\"Uneven rock surface with footprints. \" width=\"399\" height=\"245\" \/><figcaption class=\"wp-caption-text\">Figure 7.14: A few early hominin footprints fossilized at Laetoli. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:NHM_-_Laetoli_Fu%C3%9Fspuren.jpg\">NHM - Laetoli Fu\u00dfspuren<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Xenophon\">Wolfgang Sauber<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Other fossilized footprints have been discovered around the world. At Pech Merle cave in the Dordogne region of France, archaeologists discovered two fossilized footprints. They then brought in indigenous trackers from Namibia to look for other footprints. The approach worked, as many other footprints belonging to as many as five individuals were discovered with the expert eyes of the trackers (Pastoors et al. 2017). These footprints date back 12,000 years (Granger Historical Picture Archive 2018).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Some of the more unappealing but still-fascinating trace fossils are bezoars and coprolite. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_868\">Bezoars<\/a><\/strong> are hard, concrete-like substances found in the intestines of fossilized creatures. Bezoars start off like the hair balls that cats and rabbits accumulate from grooming, but they become hard, concrete-like substances in the intestines. If an animal with a hairball dies before expelling the hair ball mass <em>and <\/em>the organism becomes fossilized, that mass becomes a bezoar.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_870\">Coprolite<\/a><\/strong> is fossilized dung. One of the best collections of coprolites is affectionately known as the \u201cPoozeum.\u201d The collection includes a huge coprolite named \u201cPrecious\u201d (Figure 7.15). Coprolite, like all fossilized materials, can be <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_872\">in matrix<\/a><\/strong>\u2014meaning that the fossil is embedded in secondary rock. As unpleasant as it may seem to work with coprolites, remember that the organic material in dung has mineralized or has started to mineralize; therefore, it is no longer soft and is generally not smelly. Also, just as a doctor can tell a lot about health and diet from a stool sample, anthropologists can glean a great deal of information from coprolite about the diets of ancient animals and the environment in which the food sources existed. For instance, 65 million-year-old grass <em>phytoliths<\/em> (microscopic silica in plants) found in dinosaur coprolite in India revealed that grasses had been in existence much earlier than scientists initially believed (Taylor and O\u2019Dea 2014, 133).<\/p>\n<figure style=\"width: 312px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image19-1-1.jpg\" alt=\"Piece of fossilized poop.\" width=\"312\" height=\"224\" \/><figcaption class=\"wp-caption-text\">Figure 7.15: An extremely large coprolite named \u201cPrecious.\u201d Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Precious_the_Coprolite_Courtesy_of_the_Poozeum.jpg\">Precious the Coprolite Courtesy of the Poozeum<\/a> by <a href=\"https:\/\/poozeum.com\">Poozeum<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><em>Pseudofossils<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_874\">Pseudofossils<\/a><\/strong> are not to be mistaken for fake fossils, which have vexed scientists from time to time. A fake fossil is an item that is deliberately manipulated or manufactured to mislead scientists and the general public. In contrast, pseudofossils are not misrepresentations but rather misinterpretations of rocks that look like true fossilized remains (S. Brubaker, personal communication, March 9, 2018). Pseudofossils are the result of impressions or markings on rock, or even the way other inorganic materials react with the rock. A common example is dendrites, the crystallized deposits of black minerals that resemble plant growth (Figure 7.16). Other examples of pseudofossils are unusual or odd-shaped rocks that include various concretions and nodules. An expert can examine a potential fossil to see if there is the requisite internal structure of organic material such as bone or wood that would qualify the item as a fossil.<\/p>\n<figure style=\"width: 426px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image29.jpg\" alt=\"Rock with black branching fractal veins.\" width=\"426\" height=\"284\" \/><figcaption class=\"wp-caption-text\">Figure 7.16: A beautiful example of dendrites, a type of pseudofossil. It\u2019s easy to see how the black crystals look like plant growth. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-7\/\">Dendrites (Figure 7.25)<\/a> from the <a href=\"https:\/\/maturango.org\/\">Maturango Museum<\/a>, Ridgecrest, California, by Sarah S. King and Lee Anne Zajicek is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Dig Deeper: <del>Trace Fossils<\/del>\u00a0<em>The Power of Poop<\/em><\/h2>\n<p class=\"import-Normal\">Coprolites found in Paisley Caves, Oregon, in the United States are shedding new light on some of the earliest occupants in North America. Human coprolites are distinguished from animal coprolites through the identification of fecal biomarkers using lipids, or fats, and bile acids (Shillito et al. 2020a). Paisley Caves have 16,000 years of anthropogenic, or human-caused, deposition, with some coprolites having been dated as old as 12.8kya (Blong et al. 2020). Over 285 radiocarbon dates have been recorded from the site (Shillito et al. 2020a), making Paisley Caves one of the most well-dated archaeological sites in the United States. Coprolite analysis can be summarized in three levels, macroscopic, microscopic, and molecular. This can also be understood as analyzing the morphology (macroscopic), contents (microscopic), and residues (molecular) (Shillito et al. 2020b). Each of these levels adds a different layer of information. Coprolite shape is informative through what can be seen macroscopically, such as ingestions of basketry or cordage, small gravels and grains, and general shape. The contents of coprolites may be of the most interest to scientists because certain plants and animals can signal past environments as well as food procurement methods. Coprolites from Paisley Caves have included small pebbles and obsidian chips from butchering game, grinding plants, and general food preparation as well as small bits of fire cracked rock likely from cooking in hearths (Blong 2020). Additionally, rodent bones in coprolites included crania and vertebrae, which suggests whole consumption (Taylor et al. 2020). Insect remains are present in the coprolites as well, such as ants, Jerusalem crickets, June beetles, and darkling beetles (Blong 2020). In all, the coprolites of Paisley Caves have provided an invaluable resource to anthropologists to study the past climate and lifeways of early humans in the Americas.<\/p>\n<p class=\"import-Normal\">Coprolites can also signal past health, which is a study known as paleopathology. A study by Katelyn McDonough and colleagues (2022) focused on the identification of parasites in coprolites at Bonneville Estates Rockshelter in eastern Nevada and their link to the greater Great Basin during the Archaic, a period of time spanning 8,000\u20135,000 years ago. According to the study, parasites such as Acanthocephalans (thorny-headed worms) have been affecting the Great Basin for at least the last 10,000 years. Acanthocephalans are endoparasites, meaning parasites that live inside of their hosts. They are found worldwide and seem to have been concentrated in the Great Basin in the past. Bonneville Estates Rockshelter has been visited by humans for over 13,000 years, with parasite identification going back to nearly 7,000 years. The species identified at Bonneville Estates is <em>Moniliformis clarki<\/em>. This species parasitizes crickets and insects, a popular food source during the Archaic in the Great Basin. The parasite uses intermediate hosts to get to mammals and birds as definitive hosts. Crickets and beetles have been recorded as food materials in Paisley Caves as well. Insects have remained an important dietary staple for people of the Great Basin and are consumed raw, dried, brined, or ground into flour. Insects that remain uncooked or undercooked have a higher risk for transmission of parasites. Symptoms associated with Acanthocephalans infection are intense intestinal discomfort, anemia, and anorexia, leading to death. It is hypothesized that the consumption of basketry, cordage, and charcoal (which was also identified at Paisley Caves), sometimes associated with parasite-infected coprolites, may have been a method of treatment for the infection. Interestingly, present day infections from this parasite are rising after remaining quite rare, as detection of the parasite is occurring in insect farms.<\/p>\n<\/div>\n<h4 class=\"import-Normal\"><em>Walking to the Past<\/em><\/h4>\n<p class=\"import-Normal\">In 1974, British anthropologist Mary Leakey discovered fossilized animal tracks at Laetoli (Figure 7.17), not far from the important paleoanthropological site at Olduvai Gorge in Tanzania. A few years later, a 27-meter trail of hominin footprints were discovered at the same site. These 70 footprints, now referred to as the Laetoli Footprints, were created when early humans walked in wet volcanic ash. Before the impressions were obscured, more volcanic ash and rain fell, sealing the footprints. These series of environmental events were truly extraordinary, but they fortunately resulted in some of the most famous and revealing trace fossils ever found. Dating of the footprints indicate that they were made 3.6 mya (Smithsonian National Museum of Natural History 2018).<\/p>\n<figure style=\"width: 495px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image13-1-1.png\" alt=\"Eastern Africa map shows sites within Tanzania.\" width=\"495\" height=\"382\" \/><figcaption class=\"wp-caption-text\">Figure 7.17: Location of Laetoli site in Tanzania, Africa, with Olduvai Gorge nearby. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-7\/\">Laetoli and Olduvai Gorge sites (Figure 7.26)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Elyssa Ebding at <a href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\">GeoPlace, California State University, Chico<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Just as forensic scientists can use footprints to identify the approximate build of a potential suspect in a crime, archaeologists have read the Laetoli Footprints for clues to these early humans. The footprints clearly indicate bipedal hominins who had similar feet to those of modern humans. Analysis of the gait through computer simulation revealed that the hominins at Laetoli walked similarly to the way we walk today (Crompton 2012). More recent analyses confirm the similarity to modern humans but also indicate a gait that involved more of a flexed limb than that of modern humans (Hatala et al. 2016; Raichlen and Gordon 2017). The relatively short stride implies that these hominins had short legs\u2014unlike the longer legs of later early humans who migrated out of Africa (Smithsonian National Museum of Natural History 2018). In the context of Olduvai Gorge, where fossils of <em>Australopithecus afarensis<\/em> have been located and dated to the same timeframe as the footprints, it is likely that these newly discovered impressions were left by these same hominins.<\/p>\n<p class=\"import-Normal\">The footprints at Laetoli were made by a small group of as many as three <em>Australopithecus afarensis<\/em>, walking in close proximity, not unlike what we would see on a modern street or sidewalk. Two trails of footprints have been positively identified with the third set of prints appearing smaller and set in the tracks left by one of the larger individuals. While scientific methods have given us the ability to date the footprints and understand the body mechanics of the hominin, additional consideration of the footprints can lead to other implications. For instance, the close proximity of the individuals implies a close relationship existed between them, not unlike that of a family. Due to the size variation and the depth of impression, the footprints seem to have been made by two larger adults and possibly one child. Scientists theorize that the weight being carried by one of the larger individuals is a young child or a baby (Masao et al. 2016). Excavation continues at Laetoli today, resulting in the discovery of two more footprints in 2015, also believed to have been made by <em>Au. afarensis<\/em> (Masao et al. 2016).<\/p>\n<figure style=\"width: 482px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image10.jpg\" alt=\"Map shows Tularosa Basin.\" width=\"482\" height=\"331\" \/><figcaption class=\"wp-caption-text\">Figure 7.18: Tularosa Basin, New Mexico. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:HUC1305.jpg\">Map of Tularosa Basin<\/a> by the <a href=\"https:\/\/www.usgs.gov\/\">United States Geological Survey<\/a> is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p>But it is not just human evolution studies that can benefit from the analysis of fossil footprints. A recent discovery of fossilized footprints has rewritten what we know about the peopling of the Americas. It was originally thought that humans had been in the Americas for at least the last 15,000 years by crossing through the ice-free corridor (IFC) between the Cordilleran and Laurentide ice sheets in present-day Alaska and Canada. However, fossil footprints from the Tularosa Basin of New Mexico (see Figure 7.18) discovered in 2021 have challenged this theory. The footprints, dated between 22,860 (\u2213320) and 21,130 (\u2213250) years ago (nps.gov) based on <em>Ruppia cirrhosa <\/em>grass seeds located above and below the footprints, have shown humans have been in the Americas for much longer than previously thought. These footprints represent an adolescent individual and toddler walking through the lakebed at White Sands (see Figure 7.19), New Mexico, alongside both giant ground sloths and mammoths (Barras 2022; Wade 2021). Also present in the lakebed are footprints of camels and dire wolves (nps.gov 2022; Wade 2021).<\/p>\n<figure style=\"width: 789px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image31-1.png\" alt=\"Archaeologists on ground. Excavation with footprints. Closeups of footprints.\" width=\"789\" height=\"594\" \/><figcaption class=\"wp-caption-text\">Figure 7.19: Excavation of fossil footprints from New Mexico. Credit: <a href=\"https:\/\/www.usgs.gov\/programs\/climate-research-and-development-program\/news\/discovery-ancient-human-footprints-white\">Images of White Sands National Park Study Site Footprints<\/a> by the <a href=\"https:\/\/www.usgs.gov\/programs\/climate-research-and-development-program\">USGS Climate Research and Development Program<\/a> is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">The IFC model was upheld by a group of theorists known as \u201cClovis First,\u201d who believed the migration of people into the Americas was recent and was represented archaeologically through the Clovis projectile point toolkit. Subsequent discoveries at sites such as Cactus Hill on the east coast of the United States and Monte Verde, Chile, have demonstrated that this model wouldn\u2019t have worked. Because these sites are as old as 20,000 years and 18,500 years respectively, the IFC would have been frozen over and impassable (Gruhn 2020). Other models have been adopted to account for this, such as the coastal migration model down the west coast of North America. The more-likely migration scenario seems to be neither of these as more discoveries or antiquity continue to emerge. People may instead have migrated into the Americas before the last glacial maximum began, around 25,500\u201319,000 years ago. According to Indigenous knowledge, they have always been here. With the discovery of the White Sands footprints, it is known that humans have been in the Americas for at least 20,000 years.<\/p>\n<p class=\"import-Normal\">This discovery also reveals the importance of recognizing knowledge beyond that which is produced by the European scientific tradition. Rather than framing science in a way that runs counter to Indigenous knowledge, it can be thought that science is catching up with it. <span style=\"background-color: #00ffff;\">For instance, the Acoma Pueblo people have the word for <em>camel<\/em> in their vocabulary.<\/span> This was dismissed by scientists who assumed the word was for describing camels that were introduced to the United States in the past 100 years. However, the discovery of the White Sands footprints also included the footprints of Pleistocene camels in the same strata. Therefore, the fact that the Acoma Pueblo people have had a word for <em>camel<\/em> likely refers the Pleistocene-age megafauna camel, <em>Camelops hesternus,<\/em> rather than <em>Camelus dromedarius<\/em> or <em>Camelus bactrianus<\/em>, two present-day camel species (which are actually descendants of <em>Camelops hesternus<\/em>). Therefore, the existence of the Acoma Pueblo word for <em>camel <\/em>is not like an anomaly but rather a testament to the fact that Acoma Pueblo ancestors walked beside <em>C. hesternus<\/em> on this continent 20,000 years ago. These footprints challenge the \u201cice-free corridor\u201d expansion model, as the bridge connecting present-day Alaska and Russia into Canada would have been covered in an impenetrable ice sheet at this time. The discovery of these footprints urges scientists to reconsider further investigations at well-known Terminal Pleistocene\/Early Holocene dry lake beds in the Southwestern and Mojave deserts\u2014and to include Indigenous knowledge in their work rather than ignore it.<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Necropolitics<\/h2>\n<p class=\"import-Normal\">What are necropolitics? Necropolitics is an application of critical theory that describes how \u201cgovernments assign differential value to human life\u201d and similarly how someone is treated after they die (Verghese 2021). How is someone\u2019s death political?<\/p>\n<p class=\"import-Normal\">Consider the La Brea Woman example from the section on asphalt above. The La Brea Woman\u2019s discovery was controversial, not because she is the only person to be found in the tar pits or because of her age but also because of necropolitics. The La Brea Woman was collected in 1914 and her body was housed on display at the George C. Page Museum in Los Angeles against the wishes of the Chumash and the Tongva, two tribes whose ancestral lands include Los Angeles. The museum decided to display a skull cast instead to meet the request of the tribes which included a separate postcranial skeleton from a different individual. The updated display itself was wrought with other ethical issues, as a cast of her skull was \u201cattached to the ancient remains of a Pakistani female that was dyed dark bronze, the femurs shortened to approximate the stature of native people\u201d (Cooper 2010). In both cases, neither the individuals or their descendent communities consented to the display or grotesque modification of human remains. According to an interview conducted by LA Weekly (Cooper 2010) with Cindi Alvitre, former chair of the Gabrielino-Tongva Tribal Council, the display of Indigenous human remains is akin to voyeurism. She states \u201cIt's disheartening to me because it's very inappropriate to display any human remains. The things we do to fill the imagination of visitors. It violates human rights.\u201d It is important to listen to the wishes of Indigenous people and center their values when conducting work with their ancestors. A good source for considering places to look for archaeological research ethics before conducting fieldwork (and ideally during your research design) is the Society for American Archaeology\u2019s ethics principle list, as well as following the Indigenous Archaeology Collective.<\/p>\n<p class=\"import-Normal\">Indigenous remains are now protected in the United States due to legislation such as Native American Graves Protection and Repatriation Act (NAGPRA). You can read more about this in Chapter 15: Bioarchaeology and Forensic Anthropology. Before the passing of NAGPRA, tribes had little agency over how the bodies of their ancestors were treated by anthropologists and museums, including decisions about sampling and destructive tests. Now when archaeological field work is conducted on federal land, tribes must be consulted before work begins. This consultation process often includes what to do if human remains are encountered. Indigenous tribes are multifaceted and multivocal; each has its own rules about how to handle the remains of their ancestors. In some cases, all work on the project must be halted after the discovery of human remains. Other tribes allow for work to continue if the remains are moved and reburied. Some tribes are open to radiometric dating if it aligns with their beliefs in the afterlife. Each tribe is different, and each tribe deserves to have its wishes respected.<\/p>\n<\/div>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Voices From the Past: What Fossils Can Tell Us<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Given that so few organisms ever become fossilized, any anthropologist or fossil hunter will tell you that finding a fossil is extremely exciting. But this is just the beginning of a fantastic mystery. With the creative application of scientific methods and deductive reasoning, a great deal can be learned about the fossilized organism and the environment in which it lived, leading to enhanced understanding of the world around us.<\/p>\n<h3 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"background-color: #ccffcc;\"><strong>Dating Methods<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Context is a crucial concept in paleoanthropology and archaeology. Objects and fossils are interesting in and of themselves, but without context there is only so much we can learn from them. One of the most important contextual pieces is the dating of an object or fossil. By being able to place it in time, we can compare it more accurately with other contemporary fossils and artifacts or we can better analyze the evolution of a fossil species or artifacts. To answer the question \u201cHow do we know what we know?,\u201d you have to know how archaeologists and paleoanthropologists establish dates for artifacts, fossils, and sites.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Though accurate dating is important for context and analysis, we must consider the impact. Many of the chronometric dating methods used by anthropologists require the removal of small samples from artifacts, bones, soils, and rock. Thus these techniques are considered destructive. How much of an artifact are you willing to destroy to get your date? Sharon Clough, a Senior Environmental Officer at Cotswold Archaeology, addressed this issue in a case study from her research. She stated that \u201cthe benefit of a date did not outweigh the destruction of a valuable and finite resource\u201d (Clough 2020). The resource in question was human remains. When considering our dating options, we want to be sure that we do as little harm as possible, especially in the case of human remains (read more about this issue in the Special Topic box, \u201cNecropolitics\u201d).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Dating techniques are divided into two broad categories: relative dating methods and chronometric (sometimes called absolute) dating methods.<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"background-color: #ccffcc;\"><em>Relative Dating<\/em><\/span><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_876\">Relative dating<\/a><\/strong> methods are used first because they rely on simple observational skills. In the 1820s, Christian J\u00fcrgensen Thomsen at the National Museum of Denmark in Copenhagen developed the \u201cthree-age\u201d system still used in European archaeology today (Feder 2017, 17). He categorized the artifacts at the museum based on the idea that simpler tools and materials were most likely older than more complex tools and materials. Stone tools must predate metal tools because they do not require special technology to develop. Copper and bronze tools must predate iron because they can be smelted or worked at lower temperatures, etc. Based on these observations, he categorized the artifacts into Stone Age, Bronze Age, and Iron Age.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">The restriction of relative dating is that you don\u2019t know specific dates or how much time passed between different sites or artifacts. You simply know that one artifact or fossil is older than another. Thomsen knew that Stone Age artifacts were older than Bronze Age artifacts, but he couldn\u2019t tell if they were hundreds of years older or thousands of years older. The same is true with fossils that have differences of ages into the hundreds of millions of years.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">The first relative dating technique is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_878\">stratigraphy<\/a> <\/strong>(Figure 7.20). You might have already heard this term if you have watched documentaries on archaeological excavations. That\u2019s because this method is still being used today. It provides a solid foundation for other dating techniques and gives important context to artifacts and fossils found at a site.<\/p>\n<figure style=\"width: 382px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image11-1.png\" alt=\"Stratigraphic cross-section with 12 strata.\" width=\"382\" height=\"662\" \/><figcaption class=\"wp-caption-text\">Figure 7.20: An illustration of a stratigraphic cross-section. The objects at a lower strata are older than the one above. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-7\/\">Stratigraphic cross-section (Figure 7.28)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Stratigraphy is based on the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_880\">Law of Superposition<\/a><\/strong> first proposed by Nicholas Steno in 1669 and further explored by James Hutton (the previously mentioned \u201cFather\u201d of Deep Time). Essentially, superposition tells us that things on the bottom are older than things on the top (Williams 2004, 28). Notice on Figure 7.20 that there are distinctive layers piled on top of each other. It stands to reason that each layer is older than the one immediately on top of it (Hester et al. 1997, 338). Think of a pile of laundry on the floor. Over the course of a week, as dirty clothes get tossed on that pile, the shirt tossed down on Monday will be at the bottom of the pile while the shirt tossed down on Friday will be at the top. Assuming that the laundry pile was undisturbed throughout the week, if the clothes were picked up layer by layer, the clothing choices that week could be reconstructed in the order that they were worn.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Another relative dating technique is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_882\">biostratigraphy<\/a><\/strong>. This form of dating looks at the context of a fossil or artifact and compares it to the other fossils and biological remains (plant and animal) found in the same stratigraphic layers. For instance, if an artifact is found in the same layer as wooly mammoth remains, you know that it must date to around the last ice age, when wooly mammoths were still abundant on Earth. In the absence of more specific dating techniques, early archaeologists could prove the great antiquity of stone tools because of their association with extinct animals. The application of this relative dating technique in archaeology was used at the Folsom site in New Mexico. In 1927, a stone spear point was discovered embedded in the rib of an extinct species of bison. Because of the undeniable association between the artifact and the ancient animal, there was scientific evidence that people had occupied the North American continent since antiquity (Cook 1928).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Similar to biostratigraphic dating is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_884\">cultural dating<\/a> <\/strong>(Figure 7.21). This relative dating technique is used to identify the chronological relationships between human-made artifacts. Cultural dating is based on artifact types and styles (Hester et al. 1997, 338). For instance, a pocket knife by itself is difficult to date. However, if the same pocket knife is discovered surrounded by cassette tapes and VHS tapes, it is logical to assume that the artifact came from the late 20th century like the cassette and VHS tapes. The pocket knife could not be dated earlier than the late 20th century because the tapes were made no earlier than 1977. In the Thomsen example above, he was able to identify a relative chronology of ancient European tools based on the artifact styles, manufacturing techniques, and raw materials. Cultural dating can be used with any human-made artifacts. Both cultural dating and biostratigraphy are most effective when researchers are already familiar with the time periods for the artifacts and animals. They are still used today to identify general time periods for sites.<\/p>\n<figure style=\"width: 364px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image26-1.png\" alt=\"Ax heads, swords, circlets, and pots by type.\" width=\"364\" height=\"557\" \/><figcaption class=\"wp-caption-text\">Figure 7.21: Charts of typology, like these representing items from the Bronze Age, are used to classify artifacts and illustrate cultural material assemblages. Credit: <a href=\"https:\/\/wellcomecollection.org\/works\/de5rxx5a\">Bronze Age implements, ornaments and pottery (Period II)<\/a> by <a href=\"https:\/\/wellcomecollection.org\/\">Wellcome Collection<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/#_ga=2.5144115.1054155377.1564173886-467226638.1563307053\">CC BY 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Chemical dating was developed in the 19th century and represents one of the early attempts to use soil composition and chemistry to date artifacts. A specific type of chemical dating is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_886\">fluorine dating<\/a><\/strong>, and it is commonly used to compare the age of the soil around bone, antler, and teeth located in close proximity (Cook and Ezra-Cohn 1959; Goodrum and Olson 2009). While this technique is based on chemical dating, it only provides the relative dates of items rather than their absolute ages. For this reason, fluorine dating is considered a hybrid form of relative and chronometric dating methods (which will be discussed next).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Soils contain different amounts of chemicals, and those chemicals, such as fluorine, can be absorbed by human and animal bones buried in the soil. The longer the remains are in the soil, the more fluorine they will absorb (Cook and Ezra-Cohn 1959; Goodrum and Olson 2009). A sample of the bone or antler can be processed and measured for its fluorine content. Unfortunately, this absorption rate is highly sensitive to temperature, soil pH, and varying fluorine levels in local soil and groundwater (Goodrum and Olson 2009; Haddy and Hanson 1982). This makes it difficult to get an accurate date for the remains or to compare remains between two sites. However, this technique is particularly useful for determining whether different artifacts come from the same burial context. If they were buried in the same soil for the same length of time, their fluorine signatures would match.<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"background-color: #ccffcc;\"><em>Chronometric Dating<\/em><\/span><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Unlike relative dating methods, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_888\">chronometric dating<\/a><\/strong> methods provide specific dates and time ranges. Many of the chronometric techniques we will discuss are based on work in other disciplines such as chemistry and physics. The modern developments in studying radioactive materials are accurate and precise in establishing dates for ancient sites and remains.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Many of the chronometric dating methods are based on the measurement of radioactive decay of particular elements. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_890\">Elements<\/a><\/strong> are materials that cannot be broken down into more simple materials without losing their chemical identity (Brown et al. 2018, 48). Each element consists of an <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_892\">atom<\/a><\/strong> that has a specific number of protons (positively charged particles) and electrons (negatively charged particles) as well as varying numbers of neutrons (particles with no charge). The protons and neutrons are located in the densely compacted nucleus of the atom, but the majority of the volume of an atom is space outside the nucleus around which the electrons orbit (see Figure 7.22).<\/p>\n<figure style=\"width: 285px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image18-1-1.png\" alt=\"Atom labeled with nucleus, proton, neutron, and electron.\" width=\"285\" height=\"285\" \/><figcaption class=\"wp-caption-text\">Figure 7.22: Simplified illustration of an atom. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Atom%20Diagram.svg\">Atom Diagram<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:AG_Caesar\">AG Caesar<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Elements are classified based on the number of protons in the nucleus. For example, carbon has six protons, giving it an atomic number 6. Uranium has 92 protons, which means that it has an atomic number 92. While the number of protons in the atom of an element do not vary, the number of neutrons may. Atoms of a given element that have different numbers of neutrons are known as <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_894\">isotopes<\/a><\/strong>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">The majority of an atom\u2019s mass is determined by the protons and neutrons, which have more than a thousand times the mass of an electron. Due to the different numbers of neutrons in the nucleus, isotopes vary by nuclear\/atomic weight (Brown et al. 2018, 94). For instance, isotopes of carbon include carbon 12 (<sup>12<\/sup>C), carbon 13 (<sup>13<\/sup>C), and carbon 14 (<sup>14<\/sup>C). Carbon always has six protons, but <sup>12<\/sup>C has six neutrons whereas <sup>14<\/sup>C has eight neutrons. Because <sup>14<\/sup>C has more neutrons, it has a greater mass than <sup>12<\/sup>C (Brown et al. 2018, 95).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Most isotopes in nature are considered <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_896\">stable isotopes<\/a><\/strong> and will remain in their normal structure indefinitely. However, some isotopes are considered <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_898\">unstable isotopes<\/a><\/strong> (sometimes called radioisotopes) because they spontaneously release energy and particles, transforming into stable isotopes (Brown et al. 2018, 946; Flowers et al. 2018, section 21.1). The process of transforming the atom by spontaneously releasing energy is called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_900\">radioactive decay<\/a><\/strong>. This change occurs at a predictable rate for nearly all radioisotopes of elements, allowing scientists to use unstable isotopes to measure time passage from a few hundred to a few billion years with a large degree of accuracy and precision.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">The leading chronometric method for archaeology is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_902\">radiocarbon dating<\/a> <\/strong>(Figure 7.23). This method is based on the decay of <sup>14<\/sup>C, which is an unstable isotope of carbon. It is created when nitrogen 14 (<sup>14<\/sup>N) interacts with cosmic rays, which causes it to capture a neutron and convert to <sup>14<\/sup>C. Carbon 14 in our atmosphere is absorbed by plants during photosynthesis, a process by which light energy is turned into chemical energy to sustain life in plants, algae, and some bacteria. Plants absorb carbon dioxide from the atmosphere and use the energy from light to convert it into sugar that fuels the plant (Campbell and Reece 2005, 181\u2013200). Though <sup>14<\/sup>C is an unstable isotope, plants can use it in the same way that they use the stable isotopes of carbon.<\/p>\n<figure style=\"width: 514px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image27.png\" alt=\"Creation of Carbon 14.\" width=\"514\" height=\"658\" \/><figcaption class=\"wp-caption-text\">Figure 7.23: A graphic illustrating how 14C is created in the atmosphere, is absorbed by living organisms, and ends up in the archaeological record. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-7\/\">Radiocarbon dating (Figure 7.32)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Animals get <sup>14<\/sup>C by eating the plants. Humans take it in by eating plants and animals. After death, organisms stop taking in new carbon, and the unstable <sup>14<\/sup>C will begin to decay. Carbon 14 has a half-life of 5,730 years (Hester et al. 1997, 324). That means that in 5,730 years, half the amount of <sup>14<\/sup>C will convert back into <sup>14<\/sup>N. Because the pattern of radioactive decay is so reliable, we can use <sup>14<\/sup>C to accurately date sites up to 55,000 years old (Hajdas et al. 2021). However, <sup>14<\/sup>C can only be used on the remains of biological organisms. This includes charcoal, shell, wood, plant material, and bone. This method involves destroying a small sample of the material. Earlier methods of radiocarbon dating required at least 1 gram of material, but with the introduction of accelerator mass spectrometry (AMS), sample sizes as small as 1 milligram can now be used (Hajdas et al. 2021). This significantly reduces the destructive nature of this method.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">The use of radiocarbon dating at Denisova Cave in modern-day Russia revealed an astounding find, the first dated first-generation individual with a Neanderthal mother and Denisovan father. Vivian Slon and colleagues (2018) sequenced the genome, which revealed the individual's hybrid genetic background, and radiocarbon dated the remains, revealing the sub-adult was over 50,000 years old (Slon et al. 2018).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">As mentioned before, <sup>14<\/sup>C is unstable and ultimately decays back into <sup>14<\/sup>N. This decay is happening at a constant rate (even now, inside your own body!). However, as long as an organism is alive and taking in food, <sup>14<\/sup>C is being replenished in the body. As soon as an organism dies, it no longer takes in new <sup>14<\/sup>C. We can then use the rate of decay to measure how long it has been since the organism died (Hester et al. 1997, 324). However, the amount of <sup>14<\/sup>C in the atmosphere is not stable over time. It fluctuates based on changes to the earth\u2019s magnetic field and solar activity. In order to turn <sup>14<\/sup>C results into accurate calendar years, they must be calibrated using data from other sources. Annual tree rings (see discussion of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_904\">dendrochronology<\/a><\/strong> below), <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_906\">foraminifera<\/a><\/strong> from stratified marine sediments, and microfossils from lake sediments can be used to chart the changes in <sup>14<\/sup>C as \u201ccalibration curves.\u201d The radiocarbon date obtained from the sample is compared to the established curve and then adjusted to reflect a more accurate calendar date (see Figure 7.24). The curves are updated over time with more data so that we can continue to refine radiocarbon dates (T\u00f6rnqvist et al. 2016). The most recent calibration curves were released in 2020 and may change the dates for some existing sites by hundreds of years (Jones 2020).<\/p>\n<figure style=\"width: 547px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image17-2.jpg\" alt=\"Radiocarbon date calibration curve. \" width=\"547\" height=\"384\" \/><figcaption class=\"wp-caption-text\">Figure 7.24: This is a simplified example of a calibration curve, showing how the radiocarbon age (y axis) is compared with the calibration curve to produce calibrated dates (x axis). <a href=\"https:\/\/docs.google.com\/document\/d\/1VUDKMBJYS_jNONjLxT04jQN0_z9Ua50BRN6auGSHUuU\/edit\" target=\"_blank\" rel=\"noopener\">A full text description of this image is available<\/a>. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Radiocarbon_Date_Calibration_Curve.svg\">Radiocarbon Date Calibration Curve<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:HowardMorland\">HowardMorland<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/\">CC BY-SA 3.0 License<\/a>. [Based on information from Reimer et al. 2004. Radiocarbon 46: 1029-58.]<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">As you will see in the hominin chapters (Chapters 9\u201312), 55,000 years is only a tiny fragment of human evolutionary history. It is insignificant in the context of the age of our planet. In order to date even older fossils, other methods are necessary.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_908\">Potassium-argon (K-Ar) dating<\/a><\/strong> and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_910\">argon-argon (Ar-Ar) dating<\/a><\/strong> can reach further back into the past than radiocarbon dating. Used to date volcanic rock, these techniques are based on the decay of unstable potassium 40 (<sup>40<\/sup>K) into argon 40 (<sup>40<\/sup>Ar) gas, which gets trapped in the crystalline structures of volcanic material. It is a method of indirect dating. Instead of dating the fossil itself, K-Ar and Ar-Ar dates volcanic layers around the fossil. It will tell you when the volcanic eruption that deposited the layers occurred. This is where stratigraphy becomes important. The date of the surrounding layers can give you a minimum and maximum age of the fossil based on where it is in relation to those layers. This technique was used at Gesher Benot Ya'aqo in the Jordan Valley, dating early stratigraphic deposits of basalt flows to 100,000 years old (Bar-Yosef and Belmaker 2011). The site is unique because early layers of occupation with an Acheulean handaxe industry were made primarily of basalt, which is an uncommon material for this tool technology (see Chapter 10 for a full discussion of this tool technology). The benefit of this dating technique is that <sup>40<\/sup>K has a half-life of circa 1.3 billion years, so it can be used on sites as young as 100 kya and as old as the age of Earth. As you will see in later chapters, it is particularly useful in dating early hominin sites in Africa (Michels 1972, 120; Renfrew and Bahn 2016, 155). Another benefit to this technique is that it does not damage precious fossils because the samples are taken from the surrounding rock instead. However, this method is not without its flaws. A study by J. G. Funkhouser and colleagues (1966) and Raymond Bradley (2015) demonstrated that igneous rocks with fluid inclusions, such as those found in Hawai\u2018i, can release gasses including radiogenic argon when crushed, leading to incorrectly older dates. This is an example of why it is important to use multiple dating methods in research to detect anomalies.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_912\">Uranium series dating<\/a><\/strong> is based on the decay chain of unstable isotopes of uranium. It uses mass spectrometry to detect the ratios of uranium 238 (<sup>238<\/sup>U), uranium 234(<sup>234<\/sup>U), and thorium 230 (<sup>230<\/sup>Th) in carbonates (Wendt et al. 2021). Thorium accumulates in the carbonate sample through radiometric decay. Thus, the age of the sample is calculated from the difference between a known initial ratio and the ratio present in the sample to be dated. This makes uranium series ideal for dating carbonate rich deposits such as carbonate cements from glacial moraine deposits, speleothems (deposits of secondary minerals that form on the walls, floors, and ceilings of caves, like stalactites and stalagmites), marine and lacustrine carbonates from corals, caliche, and tufa, as well as bones and teeth (University of Arizona, n.d.; van Calsteren and Thomas 2006). Due to the timing of the decay process, this dating technique can be used from a few years up to 650k (Wendt et al. 2021). Since many early hominin sites occur in cave environments, this dating technique can be very powerful. This method has also been used to develop more accurate calibration curves for radiocarbon dating. However, the accuracy of this method depends on knowing the initial ratios of the elements and ruling out possible contamination (Wendt et al. 2021). It also involves the destruction of a small sample of material.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_914\">Fission track dating<\/a> <\/strong>is another useful dating technique for sites that are millions of years old. This is based on the decay of radioactive uranium 238 (<sup>238<\/sup>U). The unstable atom of <sup>238<\/sup>U fissions at a predictable rate. The fission takes a lot of energy and causes damage to the surrounding rock. For instance, in volcanic glasses we can see this damage as trails in the glass. Researchers in the lab take a sample of the glass and count the number of fission trails using an optical microscope. As <sup>238<\/sup>U has a half-life of 4,500 million years, it can be used to date rock and mineral material starting at just a few decades and extending back to the age of Earth. As with K-Ar, archaeologists are not dating artifacts directly. They are dating the layers around the artifacts in which they are interested (Laurenzi et al. 2007).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_916\">Luminescence dating<\/a><\/strong>, which includes thermoluminescence and a related technique called optically stimulated luminescence, is based on the naturally occurring background radiation in soils. Pottery, baked clay, and sediments that include quartz and feldspar are bombarded by radiation from the soils surrounding it. Electrons in the material get displaced from their orbit and trapped in the crystalline structure of the pottery, rock, or sediment. When a sample of the material is heated to 500\u00b0C (thermoluminescence) or exposed to particular light wavelengths (optically stimulated luminescence) in the laboratory, this energy gets released in the form of light and heat and can be measured (Cochrane et al. 2013; Renfrew and Bahn 2016, 160). You can use this method to date artifacts like pottery and burnt flint directly. When attempting to date fossils, you may use this method on the crystalline grains of quartz and feldspar in the surrounding soils (Cochrane et al. 2013). The important thing to remember with this form of dating is that heating the artifact or soils will reset the clock. The method is not necessarily dating when the object was last made or used but when it was last heated to 500\u00b0C or more (pottery) or exposed to sunlight (sediments). Luminescence dating can be used on sites from less than 100 years to over 100,000 years (Duller 2008, 4). As with all archaeological data, context is crucial to understanding the information.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Like thermoluminescence dating, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_918\">electron spin resonance dating<\/a><\/strong> is based on the measurement of accumulated background radiation from the burial environment. It is used on artifacts and rocks with crystalline structures, including tooth enamel, shell, and rock\u2014those for which thermoluminescence would not work. The radiation causes electrons to become dislodged from their normal orbit. They become trapped in the crystalline matrix and affect the electromagnetic energy of the object. This energy can be measured and used to estimate the length of time in the burial environment. This technique works well for remains as old as two million years (Carvajal et al. 2011, 115\u2013116). It has the added benefit of being nondestructive, which is an important consideration when dealing with irreplaceable material.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Not all chronometric dating methods are based on unstable isotopes and their rates of decay. There are several other methods that make use of other natural biological and geologic processes. One such method is known as dendrochronology (Figure 7.25), which is based on the natural growth patterns of trees. Trees create concentric rings as they grow; the width of those rings depends on environmental conditions and season. The age of a tree can be determined by counting its rings, which also show records of rainfall, droughts, and forest fires.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image16-1.png\" alt=\"A tree, cross-section of tree core, and tree-ring timeline.\" width=\"413\" height=\"450\" \/><\/p>\n<figure style=\"width: 411px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image23-1-1.png\" alt=\"Tree rings and dates.\" width=\"411\" height=\"424\" \/><figcaption class=\"wp-caption-text\">Figure 7.25: Dendrochronology uses the variations in tree rings to create timelines. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-7\/\">Dendrochronology (Figure 7.34)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Tree rings can be used to date wood artifacts and ecofacts from archaeological sites. This first requires the creation of a profile of trees in a particular area. The Laboratory of Tree-Ring Research at the University of Arizona has a comprehensive and ongoing catalog of tree profiles (see University of Arizona n.d.). Archaeologists can then compare wood artifacts and ecofacts with existing timelines, provided the tree rings are visible, and find where their artifacts fit in the pattern. Dendrochronology has been in use since the early 20th century (Dean 2009, 25). The Northern Hemisphere chronology stretches back nearly 14,000 years (Reimer et al. 2013, 1870) and has been used successfully to date southwestern U.S. sites such as Pueblo Bonito and Aztec Ruin (Dean 2009, 26). Dendrochronological evidence has helped calibrate radiocarbon dates and even provided direct evidence of global warming (Dean 2009, 26\u201327).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">In Australia, dendrochronology, along with other environmental reconstruction methods, has been used to show that the Indigenous people had sophisticated land management systems before the arrival of British invaders. According to the work of Michael-Shawn Fletcher and colleagues (2021), there was a significant encroachment of the rainforests and tree species into grasslands after the British invasion. Prior to this time, Indigenous people managed the landscape through controlled burns at regular intervals. This practice created climate-resistant grasslands that were biodiverse and provided predictable food supplies for humans and other animals. Under European land management, there have been negative impacts on biodiversity and climate resilience and an increase in catastrophic wildfires (Fletcher et al. 2021).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">This dating method does have its difficulties. Some issues are interrupted ring growth, microclimates, and species growth variations. This is addressed through using multiple samples, statistical analysis, and calibration with other dating methods. Despite these limitations, dendrochronology can be a powerful tool in dating archaeological sites (Hillam et al. 1990; Kuniholm and Striker 1987).<\/p>\n<h3 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Environmental Reconstruction<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">As you read in Chapter 2, Charles Darwin, Jean-Baptiste Lamarck, Alfred Russel Wallace, and others recognized the importance of the environment in shaping the evolutionary course of animal species. To understand what selective processes might be shaping evolutionary change, we must be able to reconstruct the environment in which the organism was living.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">One of the ways to do that is to look at the plant species that lived in the same time range as the species in which you are interested. One way to identify ancient flora is to analyze <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_920\">sediment cores<\/a><\/strong> from water and other protected sources. Pollen gets released into the air and some of that pollen will fall on wetlands, lakes, caves, and so forth. Eventually it sinks to the bottom of the lake and forms part of the sediment. This happens year after year, so subsequent layers of pollen build up in an area, creating strata. By taking a core sample and analyzing the pollen and other organic material, an archaeologist can build a timeline of plant types and see changes in the vegetation of the area (Hester et al. 1997, 284). This can even be done over large areas by studying ocean bed cores, which accumulate pollen and dust from large swaths of neighboring continents.<\/p>\n<p class=\"import-Normal\">While sediment coring is one of the more common ways to reconstruct past environments, there are a few other methods. These have been recently employed at Holocene Lake Ivanpah, a paleolake that straddles the California and Nevada border in the United States. This lake was originally thought to have been completely dry around 9,300\u20137,800 kya (Sims and Spaulding 2017). However, analyzing core samples using soil identification, sediment chemistry, subsurface stratigraphy, and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_922\">geomorphology<\/a><\/strong> (the study of the physical characteristics of the Earth\u2019s surface) revealed deposition of three recent lake fillings during this period in the forms of additional hardpan, or lake bottom, playas, bedded or layered fine-grained (wetland) sediments, and buried beaches below the surface (Sims and Spaulding 2017; Spaulding and Sims 2018). These discoveries are important because they have not been integrated into interpretation of the local archaeological record, as it was assumed that the lake had been dry for thousands of years. Sedimentological analyses such as coring and those listed above can provide great insight into past climates and are accomplished in a minimally destructive way.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Another way of reconstructing past environments is by using stable isotopes. Unlike unstable isotopes, stable isotopes remain constant in the environment throughout time. Plants take in the isotopes through photosynthesis and ground water absorption. Animals take in isotopes by drinking local water and eating plants. Stable isotopes can be powerful tools for identifying where an organism grew up and what kind of food the organism ate throughout its life. They can even be used to identify global temperature fluctuations.<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><em>Global Temperature Reconstruction<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Oxygen isotopes are a powerful tool in tracking global temperature fluctuations throughout time. The isotopes of Oxygen 18 (<sup>18<\/sup>O) and Oxygen 16 (<sup>16<\/sup>O) occur naturally in Earth\u2019s water. Both are stable isotopes, but <sup>18<\/sup>O has a heavier atomic weight. In the normal water cycle, evaporation takes water molecules from the surface to the atmosphere. Because <sup>16<\/sup>O is lighter, it is more likely to be part of this evaporation process. The moisture gathers in the atmosphere as clouds that eventually may produce rain or snow and release the water back to the surface of the planet. During cool periods like <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_924\">glacial periods<\/a><\/strong> (ice ages), the evaporated water often comes down to Earth\u2019s surface as snow. The snow piles up in the winter but, because of the cooler summers, does not melt off. Instead, it gets compacted and layered year after year, eventually resulting in large glaciers or ice sheets covering parts of Earth. Since <sup>16<\/sup>O, with the lighter atomic weight, is more likely to be absorbed in the evaporation process, it gets locked up in glacier formation. The waters left in oceans would have a higher ratio of <sup>18<\/sup>O during these periods of cooler global temperatures (Potts 2012, 154\u2013156; see Figure 7.26).<\/p>\n<figure style=\"width: 389px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image15-1.png\" alt=\"Graph with oxygen isotope on y axis and years on x axis.\" width=\"389\" height=\"218\" \/><figcaption class=\"wp-caption-text\">Figure 7.26: This graph depicts how temperatures of the sea have fluctuated greatly over the course of the history of the planet. <a href=\"https:\/\/docs.google.com\/document\/d\/1VUDKMBJYS_jNONjLxT04jQN0_z9Ua50BRN6auGSHUuU\/edit\" target=\"_blank\" rel=\"noopener\">A full text description of this image is available.<\/a> Credit: <a href=\"https:\/\/www.giss.nasa.gov\/research\/briefs\/1999_schmidt_01\/\">Oxygen in deep sea sediment carbonate (Figure 2)<\/a> by <a href=\"https:\/\/www.giss.nasa.gov\/\">NASA Goddard Institute for Space Studies<\/a> originally from \"Science Briefs: Cold Climates, Warm Climates: How Can We Tell Past Temperatures?\" by <a href=\"https:\/\/www.giss.nasa.gov\/staff\/gschmidt.html\">Gavin Schmidt<\/a>, is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">The microorganisms that live in the oceans, foraminifera, absorb the water from their environment and use the oxygen isotopes in their body structures. When these organisms die, they sink to the ocean floor, contributing to the layers of sediment. Scientists can extract these ocean cores and sample the remains of foraminifera for their <sup>18<\/sup>O and <sup>16<\/sup>O ratios. These ratios give us a good approximation of global temperatures deep into the past. Cooler temperatures indicate higher ratios of <sup>18<\/sup>O (Potts 2012, 154\u2013156).<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><em>Diet Reconstruction<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">You may be familiar with the saying \u201cyou are what you eat.\u201d When it comes to your teeth and bones, this adage is literal. Stable isotopes can also be used to reconstruct animal diet and migration patterns. Living organisms absorb elements from ingested plants and water. These elements are used in tissues like bones, teeth, skin, hair, and so on. By analyzing the stable isotopes in the bones and teeth of humans and other animals, we can identify the types of food they ate at different stages of their lives.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Plants take in carbon dioxide from the atmosphere during photosynthesis. We\u2019ve already discussed this using the example of the unstable isotope <sup>14<\/sup>C; however, this absorption also takes place with the stable isotopes of <sup>12<\/sup>C and <sup>13<\/sup>C. During photosynthesis, some plants incorporate carbon dioxide as a three-carbon molecule (C3 plants) and some as a four-carbon molecule (C4 plants). On the one hand, C3 plants include certain types of trees and shrubs that are found in relatively wet environments and have lower ratios of <sup>13<\/sup>C compared to <sup>12<\/sup>C. C4 plants, on the other hand, include plants from drier environments like savannahs and grasslands. C4 plants have higher ratios of <sup>13<\/sup>C to <sup>12<\/sup>C than C3 plants (Renfrew and Bahn 2016, 312). These ratios remain stable as you go up the food chain. Therefore, you can analyze the bones and teeth of an animal to identify the <sup>13<\/sup>C\/<sup>12<\/sup>C ratios and identify the types of plants that animal was eating.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">The ratios of stable nitrogen isotopes <sup>15<\/sup>N and <sup>14<\/sup>N can also give information about the diet of fossilized or deceased organisms. Though initially absorbed from water and soils by plants, the nitrogen ratios change depending on the primary diet of the organism. An animal who has a mostly vegetarian diet will have lower ratios of <sup>15<\/sup>N to <sup>14<\/sup>N, while those further up the food chain, like carnivores, will have higher ratios of <sup>15<\/sup>N. Interestingly, breastfeeding infants have a higher nitrogen ratio than their mothers, because they are getting all of their nutrients through their mother\u2019s milk. So nitrogen can be used to track life events like weaning (Jay et al. 2008, 2). A marine versus terrestrial diet will also affect the nitrogen signatures. Terrestrial diets have lower ratios of <sup>15<\/sup>N than marine diets. In the course of human evolution, this type of analysis can help us identify important changes in human nutrition. It can help anthropologists figure out when meat became a primary part of the ancient human diet or when marine resources began to be used. The ratios of stable nitrogen isotopes can also be used to determine a change in status, as in the case of the Llullaillaco children (the \u201cice mummies\u201d) found in the Andes Mountains. For instance, the nitrogen values in hair from the Llullaillaco Maiden showed a significant positive shift that is associated with increased meat consumption in the last 12 months of her life (Wilson et al. 2007). Although the two younger children had little changes in their diets in the last year of their short lives, the changes in their nitrogen values were significant enough to suggest that the improvement in their diets may have been attributed to the Incas\u2019 desire to sacrifice healthy, high-status children\u201d (Faux 2012, 6).<\/p>\n<h4 class=\"import-Normal\"><em>Migration<\/em><\/h4>\n<p class=\"import-Normal\">Stable isotopes can also tell us a great deal about where an individual lived and whether they migrated during their lifetime. The geology of Earth varies because rocks and soils have different amounts or ratios of certain elements in them. These variations in the ratios of isotopes of certain elements are called isotopic signatures. They are like a chemical fingerprint for a geographical region. These isotopes get into the groundwater and are absorbed by plants and animals living in that area. Elements like strontium, oxygen, and nitrogen, among others, are then used by the body to build bones and teeth. If you ate and drank local water all of your life, your bones and teeth would have the same isotopic signature as the geographical region in which you lived.<\/p>\n<p class=\"import-Normal\">However, many people (and animals) move around during their lifetimes. Isotopic signatures can be used to identify migration patterns in organisms (Montgomery et al. 2005). Teeth develop in early childhood. If the isotopes of teeth are analyzed, these isotopes would resemble those found in the geographic area where an individual lived as a child. Bones, however, are a different story. Bones are constantly changing throughout life. Old cells are removed and new cells are deposited to respond to growth, healing, activity change, and general deterioration. Therefore, the isotopic signature of bones will reflect the geographical area in which an individual spent the last seven to ten years of life. If an individual has different isotopic signatures for their bones and teeth, it could indicate a migration some time during their life after childhood.<\/p>\n<figure style=\"width: 386px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image24-2.jpg\" alt=\"Upright boulders of Stonehenge.\" width=\"386\" height=\"289\" \/><figcaption class=\"wp-caption-text\">Figure 7.27: Stonehenge continues to provide clues to its mysterious existence with recent research using isotope ratios. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-7\/\">Stonehenge (Figure 7.37)<\/a> by Sarah S. King is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p>Recent work involving stable isotope analysis has been done on the cremation burials from Stonehenge, in Wessex, England (Figure 7.27). Much of the archaeological work at Stonehenge in the past focused on the building and development of the monument itself. That is partly because most of the burials at the monument were cremated remains, which are difficult to study because of their fragmentary nature and the chemical alterations that bone and teeth undergo when heated. The cremation process complicates the oxygen and carbon isotopes. However, the researchers determined that strontium would not be affected by heating and could still be analyzed in cranial fragments. Using the remains of 25 individuals, they compared their strontium signatures to the geology of Wessex and other regions of the UK. Fifteen of those individuals had strontium signatures that matched the local geology. This means that in the last ten or so years of their lives, they lived and ate food from around Stonehenge. However, ten of the individuals did not match the local geologic signature. These individuals had strontium ratios more closely aligned with the geology of west Wales. Archaeologists find this particularly interesting because in the early phases of Stonehenge\u2019s construction, the smaller \u201cblue stones\u201d were brought 200 km from Wales in a feat of early engineering. These larger regional connections show that Stonehenge was not just a site of local importance. It dominated a much larger region of influence and drew people from all over ancient Britain (Snoeck et al. 2018).<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Cold Case Naia<\/h2>\n<figure style=\"width: 455px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image4-1-2.png\" alt=\"Sites on Yucatan peninsula.\" width=\"455\" height=\"351\" \/><figcaption class=\"wp-caption-text\">Figure 7.28: Map of Mexico showing the Yucatan Peninsula and the locations of Hoyo Negro and Sistema Sac Actun. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-7\/\">Hoyo Negro and Sistema Sac Actun, Mexic0 (Figure 7.38)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Elyssa Ebding at <a href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\">GeoPlace, California State University, Chico<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">In 2007, cave divers exploring the Sistema Sac Actun in the Yucat\u00e1n Peninsula in Mexico (see Figure 7.28 and 7.29) discovered the bones of a 15- to 16-year-old female human along with the bones of various extinct animals from the Pleistocene (Collins et al. 2015). The site was named Hoyo Negro (\u201cBlack Hole\u201d). The human bones belonged to a Paleo-American, later named \u201cNaia\u201d after a Greek water nymph. Examination of the partially fossilized remains revealed a great deal about Naia\u2019s life, and the radiocarbon dating of her tooth enamel indicated that she lived some 13,000 years ago (Chatters et al. 2014). Naia\u2019s arms were not overly developed, so her daily activities did not involve heavy carrying or grinding of grain or seeds. Her legs, however, were quite muscular, implying that Naia was used to walking long distances. Naia\u2019s teeth and bones indicate habitually poor nutrition. There is evidence of violent injury during the course of Naia\u2019s life from a healed spiral fracture of her left forearm. Naia also suffered from tooth decay and osteoporosis even though she appeared young and undersized. Dr. Jim Chatters hypothesizes that Naia entered the cave at a time when it was not flooded, probably looking for water. She may have become disoriented and fell off a high ledge to her death. The trauma to her pelvis is consistent with such an injury (Watson 2017).<\/p>\n<p class=\"import-Normal\">Naia\u2019s skeleton is remarkably complete given its age. As divers were able to locate her skull, Naia\u2019s physical appearance in life could be interpreted. Surprisingly, in examining the skull, it was determined that Naia did not resemble modern Indigenous peoples in the region. However, the<strong> <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_926\">mitochondrial DNA<\/a><\/strong> (mtDNA) recovered from a tooth indicates that Naia shares her DNA with modern Indigenous peoples (Chatters et al. 2014). Though Naia\u2019s burial environment made chemical analysis difficult, researchers were able to recover carbon isotopes from her remains. The isotopes from Naia\u2019s tooth enamel suggest a diet of \u201ccool-season grasses and\/or broad-leaf vegetation\u201d (Chatters et al. 2022, 68). Naia\u2019s teeth also displayed numerous dental caries and only light dental wear. Coupled with the isotopic data, she likely had a \u201csofter, more sugar-rich diet\u201d (Chatters et al. 2022, 68).<\/p>\n<figure style=\"width: 625px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image32-1.png\" alt=\"Cross-section of the Hoyo Negro cenote.\" width=\"625\" height=\"353\" \/><figcaption class=\"wp-caption-text\">Figure 7.29: A diagram of the Sistema Sac Actun and the Hoyo Negro cenote where Naia rested underwater for roughly 13,000 years. The illustration depicts a cenote or hole in the ground leading to a long, narrow tunnel, ending in a large cavern. The cavern and tunnel are both filled with water. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-7\/\">Hoyo Negro cenote (Figure 7.39)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<\/div>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"background-color: #ff99cc;\">Summary<\/span><\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"background-color: #ff99cc;\">With a timeline that extends back some 4.6 billion years, Earth has witnessed continental drift, environmental changes, and a growing complexity of life. Fossils, the mineralized remains of living organisms, provide physical evidence of life and the environment on the planet over the course of billions of years. In order to better understand the fossil record, anthropologists rely on the collaboration of numerous academic fields and disciplines. Anthropologists use a variety of scientific methods, both relative and chronometric, to analyze fossils to determine age, origins, and migration patterns as well as to provide insight into the health and diet of the fossilized organism. While each method has its advantages, disadvantages, and limited applications, these tools enable anthropologists to theorize how all living organisms evolved, including the evolution of early humans into modern humans, <em>H. sapiens<\/em>. The fossil record is far from complete, but our expanding understanding of the fossil context, with exciting new discoveries and improved scientific methods, enables us to document the history of our planet and the evolution of life on Earth.<\/span><\/p>\n<h3 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Dating Methods Quick Guide<\/strong><\/h3>\n<div style=\"text-align: left;\">\n<table style=\"width: 463.5pt; height: 586px;\">\n<thead>\n<tr style=\"height: 24.25pt;\">\n<td class=\"Table1-C\" style=\"padding: 5pt; border: 1pt solid #000000; height: 30px; width: 157.45px;\">\n<p class=\"import-Normal\"><strong>Method<\/strong><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 1pt 1pt 1pt 0.75pt; padding: 5pt; height: 30px; width: 250.533px;\">\n<p class=\"import-Normal\"><strong>Material <\/strong><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 1pt 1pt 1pt 0.75pt; padding: 5pt; height: 30px; width: 166.017px;\">\n<p class=\"import-Normal\"><strong>Effective date range<\/strong><\/p>\n<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table1-R\" style=\"height: 24.25pt;\">\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt; padding: 5pt; height: 30px; width: 157.45px;\">\n<p class=\"import-Normal\">Stratigraphy<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 30px; width: 250.533px;\">\n<p class=\"import-Normal\">Soil layers<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 30px; width: 166.017px;\">\n<p class=\"import-Normal\">Relative<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 37.75pt;\">\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt; padding: 5pt; height: 36px; width: 157.45px;\">\n<p class=\"import-Normal\">Biostratigraphy<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 36px; width: 250.533px;\">\n<p class=\"import-Normal\">Plant and animal remains<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 36px; width: 166.017px;\">\n<p class=\"import-Normal\">Relative<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 24.25pt;\">\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt; padding: 5pt; height: 30px; width: 157.45px;\">\n<p class=\"import-Normal\">Cultural dating<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 30px; width: 250.533px;\">\n<p class=\"import-Normal\">Human-made objects<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 30px; width: 166.017px;\">\n<p class=\"import-Normal\">Relative<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 24.25pt;\">\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt; padding: 5pt; height: 30px; width: 157.45px;\">\n<p class=\"import-Normal\">Fluorine<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 30px; width: 250.533px;\">\n<p class=\"import-Normal\">Bone, antler, teeth<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 30px; width: 166.017px;\">\n<p class=\"import-Normal\">Relative<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 78.25pt;\">\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt; padding: 5pt; height: 90px; width: 157.45px;\">\n<p class=\"import-Normal\">Radiocarbon<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 90px; width: 250.533px;\">\n<p class=\"import-Normal\">Organic carbon bearing material (bones, teeth, antler, plant material, shell, charcoal)<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 90px; width: 166.017px;\">\n<p class=\"import-Normal\">Younger than 55,000 years<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 37.75pt;\">\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt; padding: 5pt; height: 46px; width: 157.45px;\">\n<p class=\"import-Normal\">Potassium-argon and argon-argon<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 46px; width: 250.533px;\">\n<p class=\"import-Normal\">Volcanic rock<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 46px; width: 166.017px;\">\n<p class=\"import-Normal\">Older than 100,000 years<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 64.75pt;\">\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt; padding: 5pt; height: 72px; width: 157.45px;\">\n<p class=\"import-Normal\">Uranium series<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 72px; width: 250.533px;\">\n<p class=\"import-Normal\">Carbonates such as stalactites, stalagmites, corals, caliche, and tufa<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 72px; width: 166.017px;\">\n<p class=\"import-Normal\">Younger than 650,000 years<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 37.75pt;\">\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt; padding: 5pt; height: 46px; width: 157.45px;\">\n<p class=\"import-Normal\">Fission track<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 46px; width: 250.533px;\">\n<p class=\"import-Normal\">Volcanic glasses and crystalline minerals<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 46px; width: 166.017px;\">\n<p class=\"import-Normal\">Spans age of Earth<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 37.75pt;\">\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt; padding: 5pt; height: 46px; width: 157.45px;\">\n<p class=\"import-Normal\">Luminescence<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 46px; width: 250.533px;\">\n<p class=\"import-Normal\">Pottery, baked clay, sediments<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 46px; width: 166.017px;\">\n<p class=\"import-Normal\">100 to older than 100,000 years<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 51.25pt;\">\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt; padding: 5pt; height: 54px; width: 157.45px;\">\n<p class=\"import-Normal\">Electron spin resonance dating<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 54px; width: 250.533px;\">\n<p class=\"import-Normal\">Tooth enamel, shell, rock with crystalline structures<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 54px; width: 166.017px;\">\n<p class=\"import-Normal\">Younger than 2 million years<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 51.25pt;\">\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt; padding: 5pt; height: 61px; width: 157.45px;\">\n<p class=\"import-Normal\">Dendrochronology<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 61px; width: 250.533px;\">\n<p class=\"import-Normal\">Wood (where tree rings are identifiable)<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-color: #000000; border-style: solid; border-width: 0.75pt 1pt 1pt 0.75pt; padding: 5pt; height: 61px; width: 166.017px;\">\n<p class=\"import-Normal\">Dependent on location and available chronologies<\/p>\n<\/td>\n<\/tr>\n<tr style=\"height: 15px;\">\n<td style=\"height: 15px; width: 158.317px;\"><\/td>\n<td style=\"height: 15px; width: 251.9px;\"><\/td>\n<td style=\"height: 15px; width: 166.883px;\"><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Review Questions<\/h2>\n<ul>\n<li class=\"import-Normal\" style=\"text-indent: 0pt;\">How do remains become fossils? What conditions are necessary for the fossilization process?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt;\">What kind of information could you acquire from a single fossil? What could it tell you about the broader environment?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt;\">What factors would you take into consideration when deciding which dating method to use for a particular artifact?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt;\">What methods do anthropologists use to reconstruct past environments and lifestyles?<\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Key Terms<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Anaerobic<\/strong>: An oxygen-free environment.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Anthropocene<\/strong>: The proposed name for our current geologic epoch based on human-driven climate change.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Argon-argon (Ar-Ar) dating<\/strong>: A chronometric dating method that measures the ratio of argon gas in volcanic rock to estimate time elapsed since the volcanic rock cooled and solidified. See also <em>potassium-argon dating<\/em>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Atom<\/strong>: A small building block of matter.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Bezoars<\/strong>: Hard, concrete-like substances found in the intestines of fossil creatures.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Biostratigraphy<\/strong>: A relative dating method that uses other plant and animal remains occurring in the stratigraphic context to establish time depth.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Bya<\/strong>: Billion years ago.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Chronometric dating<\/strong>: Dating methods that give estimated numbers of years for artifacts and sites.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Continental drift<\/strong>: The slow movement of continents over time.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Coprolite<\/strong>: Fossilized poop.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Cultural dating<\/strong>: The relative dating method that arranges human-made artifacts in a time frame from oldest to youngest based on material, production technique, style, and other features.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Deep Time<\/strong>: James Hutton\u2019s theory that the world was much older than biblical explanations allowed. This age could be determined by gradual natural processes like soil erosion.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Dendrochronology<\/strong>: A chronometric dating method that uses the annual growth of trees to build a timeline into the past.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Electron spin resonance dating<\/strong>: A chronometric dating method that measures the background radiation accumulated in material over time.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Element<\/strong>: Matter that cannot be broken down into smaller matter.<\/p>\n<p class=\"import-Normal\"><strong>Eon<\/strong>: The largest unit of geologic time, spanning billions of years and divided into subunits called <em>eras<\/em>, <em>periods<\/em>, and <em>epochs<\/em>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Epochs<\/strong>: The smallest units of geologic time, spanning thousands to millions of years.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Eras<\/strong>: Units of geologic time that span millions to billions of years and that are subdivided into <em>periods<\/em> and <em>epochs<\/em>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Fission track dating<\/strong>: A chronometric dating method that is based on the fission of <sup>283<\/sup>U.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Fluorine dating<\/strong>: A relative dating method that analyzes the absorption of fluorine in bones from the surrounding soils.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Foraminifera<\/strong>: Single-celled marine organisms with shells.<\/p>\n<p class=\"import-Normal\"><strong>Fossilization<\/strong>: The process by which an organism becomes a fossil.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Fossils<\/strong>: Mineralized copies of organisms or activity imprints.<\/p>\n<p class=\"import-Normal\"><strong>G<\/strong><strong>eomorphology<\/strong>: The study of the physical characteristics of the Earth\u2019s surface.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Glacial periods<\/strong>: Periods characterized by low global temperatures and the expansion of ice sheets on Earth\u2019s surface.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Holocene<\/strong>: The geologic epoch from 10 kya to present. (See the discussion on \u201cthe Anthropocene\u201d for the debate regarding the current epoch name.)<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Hominin<\/strong>: The term used for humans and their ancestors after the split with chimpanzees and bonobos.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>In matrix<\/strong>: When a fossil is embedded in a substance, such as igneous rock.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Isotopes<\/strong>: Variants of elements.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Kya<\/strong>: Thousand years ago.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Law of Superposition<\/strong>: The scientific law that states that rock and soil are deposited in layers, with the youngest layers on top and the oldest layers on the bottom.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Lithification<\/strong>: The process by which the pressure of sediments squeeze extra water out of decaying remains and replace the voids that appear with minerals from the surrounding soil and groundwater.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Luminescence dating<\/strong>: The chronometric dating method based on the buildup of background radiation in pottery, clay, and soils.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Megafauna<\/strong>: Large animals such as mammoths and mastodons.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Mitochondrial DNA<\/strong>: DNA located in the mitochondria of a cell that is only passed down from biological mother to child.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Mya<\/strong>: Million years ago.<\/p>\n<p class=\"import-Normal\"><strong>P<\/strong><strong>aleopathology<\/strong>: Study of ancient diseases and injuries identified through examining remains.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Periods<\/strong>: Geologic time units that span millions of years and are subdivided into <em>epochs<\/em>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Permineralization<\/strong>: When minerals from water impregnate or replace organic remains, leaving a fossilized copy of the organism.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Petrified wood<\/strong>: A fossilized piece of wood in which the original organism is completely replaced by minerals through petrifaction.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Potassium-argon (K-Ar) dating<\/strong>: A chronometric dating method that measures the ratio of argon gas in volcanic rock to estimate time elapsed since the volcanic rock cooled and solidified. See also <em>argon-argon dating<\/em>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Pseudofossils<\/strong>: Natural rocks or mineral formations that can be mistaken for fossils.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Radioactive decay<\/strong>: The process of transforming the atom by spontaneously releasing energy.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Radiocarbon dating<\/strong>: The chronometric dating method based on the radioactive decay of <sup>14<\/sup>C in organic remains.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Relative dating<\/strong>: Dating methods that do not result in numbers of years but, rather, in relative timelines wherein some organisms or artifacts are older or younger than others.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Sediment cores<\/strong>: Core samples taken from lake beds or other water sources for analysis of their pollen.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Stable isotopes<\/strong>: Variants of elements that do not change over time without outside interference.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Stratigraphy<\/strong>: A relative dating method that is based on ordered layers or (strata) that build up over time.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Taphonomy<\/strong>: The study of what happens to an organism after death.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Trace fossils<\/strong>: Fossilized remains of activity such as footprints.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Uniformitarianism<\/strong>: The theoretical perspective that the geologic processes observed today are the same as the processes operating in the past.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Unstable isotopes<\/strong>: Variants of elements that spontaneously change into stable isotopes over time.<\/p>\n<p class=\"import-Normal\"><strong>Uranium series dating<\/strong>: A radiometric dating method based on the decay chain of unstable isotopes of <sup>238<\/sup>U and <sup>235<\/sup>U.<\/p>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">About the Authors<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image9-1.jpg\" alt=\"A woman with curly blonde hair and sunglasses smiles in front of a mountain landscape.\" width=\"179\" height=\"250\" \/><\/p>\n<h3 class=\"import-Normal\">Sarah S. King, Ph.D.<\/h3>\n<p class=\"import-Normal\">Cerro Coso Community College, sarah.king1@cerrocoso.edu<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Dr. Sarah S. King is an anthropology\/sociology professor at Cerro Coso Community College in California. She completed her Ph.D. work at the Division of Archaeological, Geographical and Environmental Sciences at the University of Bradford in West Yorkshire, England. Her thesis was entitled \u201cWhat Makes War?: Assessing Iron Age Warfare through Mortuary Behavior and Osteological Patterns of Violence.\u201d She also holds anthropology degrees from the University of California, Santa Cruz (B.A. hons., 2004), and the University of New Mexico, Albuquerque (M.A., 2006).<\/p>\n<p>&nbsp;<\/p>\n<h3 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><img class=\"alignleft wp-image-253\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image3-e1684810044390.jpg\" alt=\"A person with short, curly brown hair and glasses smiles at the camera.\" width=\"180\" height=\"254\" \/>Kara Jones, M.A.<\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">PhD student at University of Nevada, Las Vegas, jonesk44@unlv.nevada.edu<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Kara Jones received their B.A. in anthropology at California State University, Bakersfield (2018) and their M.A. from University of Nevada, Las Vegas (2023, summer). Their master\u2019s thesis is titled \u201cRockin\u2019 at the Lake: Toolstone Use and Procurement along Holocene Lake Ivanpah, CA.\u201d Mx Jones is a Mojave Desert archaeologist specializing in stone tool use and manufacture, focusing further on Holocene lakeshore adaptations.<\/p>\n<\/div>\n<h2>For Further Exploration<\/h2>\n<div class=\"__UNKNOWN__\">\n<h3 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Books<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Bjornerud, Marcia. 2006. <em>Reading the Rocks: The Autobiography of the Earth<\/em>. New York: Basic Books.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Hazen, Robert M. 2013. <em>The Story of Earth: The First 4.5 Billion Years, From Stardust to Living Planet<\/em>. New York: Viking Penguin.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Holmes, Richard. 2010. <em>The Age of Wonder: The Romantic Generation and the Discovery of the Beauty and Terror of Science<\/em>. New York: Vintage.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Palmer, Douglas. 2005. <em>Earth Time: Exploring the Deep Past from Victorian England to the Grand Canyon<\/em>. New York: John Wiley &amp; Sons.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Prothero, Donald R. 2015. <em>The Story of Life in 25 Fossils: Tales of Intrepid Fossil Hunters and the Wonder of Evolution<\/em>. New York: Columbia University Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Pyne, Lydia. 2016. <em>Seven Skeletons: The Evolution of the World\u2019s Most Famous Human Fossils<\/em>. New York: Viking Books.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Repcheck, Jack. 2009. <em>The Man Who Found Time: James Hutton and the Discovery of the Earth\u2019s Antiquity<\/em>. New York: Basic Books.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Taylor, Paul D., Aaron O\u2019Dea. 2014. <em>A History of Life in 100 Fossils<\/em>. Washington, DC: Smithsonian Books.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Ward, David. 2002. <em>Smithsonian Handbooks: Fossils<\/em>. Washington, DC: Smithsonian Books.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Winchester, Simon. 2009. <em>The Map That Changed the World: William Smith and the Birth of Modern Geology<\/em>. New York: Harper Perennial.<\/p>\n<h3 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><strong>Websites<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><a href=\"https:\/\/www.ambermuseum.eu\/en\/\">Amber Museum<\/a><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><a href=\"https:\/\/www.etsu.edu\/cas\/paleontology\/\">East Tennessee State University Center of Excellence in Paleontology<\/a><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><a href=\"https:\/\/www.granger.com\/\">Granger Historical Picture Archive<\/a><\/p>\n<p class=\"import-Normal\"><a href=\"https:\/\/www.facebook.com\/indigarchs\/\">Indigenous Archaeology Collective<\/a><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><a href=\"https:\/\/tarpits.org\">La Brea Tar Pits Museum<\/a><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><a href=\"https:\/\/www.lymeregismuseum.co.uk\">Lyme Regis Museum<\/a><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><a href=\"https:\/\/www.nhm.ac.uk\/discover\/mary-anning-unsung-hero.html\">Natural History Museum (London), on Mary Anning<\/a><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><a href=\"https:\/\/en.pechmerle.com\">Pech Merle Cave<\/a><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><a href=\"https:\/\/www.nps.gov\/pefo\/index.htm\">Petrified Forest National Park (NE Arizona)<\/a><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><a href=\"https:\/\/poozeum.com\">Poozeum: The No. 2 Wonder of the World<\/a><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><a href=\"https:\/\/paleobiology.si.edu\/fossiLab\/projects.html\">Smithsonian National Museum of Natural History, Department of Paleobiology<\/a><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Smithsonian National Museum of Natural History, on <a href=\"https:\/\/humanorigins.si.edu\">\u201cWhat Does It Mean to be Human\u201d<\/a><\/p>\n<p class=\"import-Normal\">Society for American Archaeology, on <a href=\"https:\/\/www.saa.org\/career-practice\/ethics-in-professional-archaeology\">\u201cEthics in Professional Archaeology\u201d<\/a><\/p>\n<p class=\"import-Normal\">Society for American Archaeology, <a href=\"https:\/\/archaeologicalethics.org\/code-of-ethics\/society-for-american-archaeology-principles-of-archaeological-ethics\/\">\u201cPrinciples of Archaeological Ethics\u201d<\/a><\/p>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">References<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Antoine, Pierre-Oliver, Maeva J. Orliac, Gokhan Atici, Inan Ulusoy, Erdal Sen, H. Evren \u00c7ubuk\u00e7u, Ebru lbayrak, Ne\u015fe Oyal, Erkan Aydar, and Sevket Sen. 2012. \u201cA Rhinocerotid Skull Cooked to Death in a 9.2 Mya-Old Ignimbrite Flow of Turkey.\u201d <em>PLoS ONE<\/em> 7 (11): e49997.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Aufderheide, Arthur C. 2003. <em>The Scientific Study of Mummies<\/em>. Cambridge, UK: Cambridge University Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Bar-Yosef, O., and M. Belmaker. 2011. \u201cEarly and Middle Pleistocene Faunal and Hominins Dispersals through Southwestern Asia.\u201d<em> Quaternary Science Reviews<\/em> 30 (11\u201312): 1318\u20131337.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Barras, C. 2022. \u201cLost Footprints of Our Ancestors.\u201d <em>New Scientist<\/em> 254 (3381): 40\u201344.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Blong, John C., Martin E. Adams, Gabriel Sanchez, Dennis L. Jenkins, Ian D. Bull, and Lisa-Marie Shillito. 2020. \u201cYounger Dryas and Early Holocene Subsistence in the Northern Great Basin: Multiproxy Analysis of Coprolites from the Paisley Caves, Oregon, USA.\u201d <em>Archaeological and Anthropological Sciences<\/em> 12 (9): 1\u201329.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Boaz, Noel T., Russel L. Ciochon, Qinqi Xu, and Jinyi Liu. 2004. \u201cMapping and Taphonomic Analysis of the <em>Homo erectus<\/em> Loci at Locality 1 Zhoukoudian, China.\u201d <em>Journal of Human Evolution <\/em>46 (5): 519\u2013549.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Booth, Thomas J., Andrew T. Chamberlain, and Mike Parker Pearson. 2015. \u201cMummification in Bronze Age Britain.\u201d <em>Antiquity<\/em> 89 (347): 1,155\u20131,173.<\/p>\n<p class=\"import-Normal\">Bradley, Raymond S. 2015. \u201cChapter 3: Dating Methods I.\u201d In <em>Paleoclimatology<\/em>, edited by Raymond S. Bradley, 55\u2013101. Cambridge, MA: Academic Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Brown, Theodore L., H. Eugene LeMay Jr., Bruce E. Burston, Catherine J. Murphy, Patrick M. Woodward, and Matthew Stoltzfus. 2018. <em>Chemistry: The Central Science.<\/em> New York: Pearson.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Campbell, Neil A., and Jane B. Reece. 2005. <em>Biology 7th ed. <\/em>New York: Pearson.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Carvajal, Eduar, Luis Montes, and Ovidio A. Almanza. 2011. \u201cQuaternary Dating by Electron Spin Resonance (ESR) Applied to Human Tooth Enamel.\u201d <em>Earth Sciences Research Journal<\/em> 15 (2): 115\u2013120.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Chatters, James C., Joaquin Arroyo-Cabrales, and Pilar Luna-Erreguerena. 2022. \u201cThe Pre-Ceramic Skeletal Record of Mexico and Central America.\u201d In <em>The Routledge Handbook of Mesoamerican Bioarchaeology,<\/em> edited by V. Tieslar, 49\u201374. New York: Routledge.<\/p>\n<p class=\"import-Normal\">Chatters, James C., Douglas J. Kennett, Yemane Asmerom, Brian M. Kemp, Victor Polyak, Alberto Nava Blank, Patricia A. Beddows, et al. 2014. \u201cLate Pleistocene Human Skeleton and mtDNA Link Paleoamericans and Modern Native Americans.\u201d <em>Science<\/em> 344 (6185): 750\u2013754.<\/p>\n<p class=\"import-Normal\">Clough, Sharon. 2020. \"Ethics in Human Osteology.\" <em>The Archaeologist<\/em> 109 (2020): 3\u20135.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Cochrane, Grant W. G., Trudy Doelman, and Lyn Wadley. 2013. \u201cAnother Dating Revolution for Prehistoric Archaeology?\u201d <em>Journal of Archaeological Method and Theory<\/em> 20 (1): 42\u201360.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Collins, S. V., E. G. Reinhardt, D. Rissolo, J. C. Chatters, A. Nava-Blank, and P. Luna-Erreguerena. 2015. \u201cReconstructing Water Level in Hoyo Negro, Quintana Roo, Mexico: Implications for Early Paleoamerican and Faunal Access.\u201d <em>Quaternary Science Reviews <\/em>124: 68\u201383.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Cook, Harold J. 1928. \u201cGlacial Age Man in New Mexico.\u201d <em>Scientific American<\/em> 139 (1): 38\u201340.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Cook, S. F., and H. C. Ezra-Cohn. 1959. \u201cAn Evaluation of the Fluorine Dating Method.\u201d <em>Southwestern Journal of Anthropology <\/em>15 (3): 276\u2013290.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Cooper, Arnie. 2010. \u201cSticky Situation at the Tar Pits.\u201d <em>LA Weekly<\/em>, May 27, 2010. https:\/\/www.laweekly.com\/sticky-situation-at-the-tar-pits\/.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Crompton, Robin H., Todd C. Pataky, Russell Savage, Kristiaan D\u2019Ao\u00fbt, Matthew R. Bennett, Michael H. Day, Karl Bates, Sarita Morse, and William I. Sellers. 2012. \u201cHuman-like External Function of the Foot, and Fully Upright Gait, Confirmed in the 3.66 Million Year Old Laetoli Hominin Footprints by Topographic Statistics, Experimental Footprint-Formation and Computer Simulation.\u201d <em>Journal of the Royal Society Interface<\/em> 9 (69): 707\u2013719. doi: 10.1098\/rsif.2011.0258<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Crutzen, Paul J., and Eugene F. Stoermer. 2000. \u201cThe \u2018Anthropocene.\u2019\u201d <em>Global Change Newsletter<\/em> 41: 17\u201318.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Darwin, Charles. 1859. <em>On the Origin of Species<\/em>. London, UK: John Murray.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Dean, Jeffery S. 2009. \u201cOne Hundred Years of Dendroarchaeology: Dating, Human Behavior, and Past Climate.\u201d In <em>Tree-rings, Kings, and Old World Archaeology and Environment: Papers Presented in Honor of Peter Ian Kuniholm<\/em>, edited by S. Manning and M. J. Bruce, 25\u201332. Oxford, UK: Oxbow Books.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Dolnick, Edward. 2011. <em>The Clockwork Universe: Isaac Newton, the Royal Society, and the Birth of the Modern World<\/em>. New York: HarperCollins.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Duller, G.A.T. 2008. <em>Luminescence Dating: Guidelines on Using Luminescence Dating in Archaeology<\/em>. Swindon, UK: English Heritage.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Eisenbeiss, Sabine. 2016. \u201cPreserved in Peat: Decoding Bodies from Lower Saxony, Germany.\u201d <em>Expedition<\/em> 58 (2): 18\u201321.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Emling, Shelley. 2009. <em>The Fossil Hunter: Dinosaurs, Evolution, and the Woman Whose Discoveries Changed the World<\/em>. New York: St. Martin\u2019s Griffin.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Faux, Jennifer L. 2012. \u201cHail the Conquering Gods: Ritual Sacrifice of Children in Inca Society.\u201d <em>Journal of Contemporary Anthropology<\/em> 3 (1): 1\u201315.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Feder, Kenneth L. 2017. <em>The Past in Perspective: An Introduction to Human Prehistory<\/em>. 7th ed. New York: Oxford University Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Fletcher, Michael-Shawn, Tegan Hall, and Andreas Nicholas Alexandra. 2021. \u201cThe Loss of an Indigenous Constructed Landscape Following British Invasion of Australia: An Insight into the Deep Human Imprint on the Australian Landscape.\u201d <em>Ambio<\/em> 50 (1): 138\u2013149.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Flowers, Paul, Klaus Theopold, Richard Langley, and William R. Robinson. 2018. <em>Chemistry<\/em>. Houston, TX: Openstax, Rice University.<\/p>\n<p class=\"import-Normal\">Fumagalli, Matteo, Ida Moltke, Niels Grarup, Fernando Racimo, Peter Bjerregaard, Marit E. J\u00f8rgensen, Thorfinn S. Korneliussen, Pascale Gerbault, Line Skotte, Allan Linneberg, Cramer Christensen, Ivan Brandslund, Torben J\u00f8rgensen, Emilia Huerta-S\u00e1nchez, Erik B. Schmidt, Oluf Pedersen, Torben Hansen, Anders Albrechtsen, and Rasmus Nielsen. 2015. \u201cGreenlandic Inuit show genetic signatures of diet and climate adaptation.\u201d <em>Science<\/em> 349 (6254): 1343-1347.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Funkhouser, J. G., I. L. Barnes, and J. J. Naughton. 1966. \u201dProblems in the Dating of Volcanic Rocks by the Potassium-Argon Method.\u201d<em> Bull Volcanol<\/em> 29: 709\u2013717.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Giles, Melanie. 2020. <em>Bog Bodies: Face to Face with the Past.<\/em> Manchester, UK: Manchester University Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Goodrum, Matthew R., and Cora Olson. 2009. \u201cThe Quest for an Absolute Chronology in Human Prehistory: Anthropologists, Chemists, and the Fluorine Dating Method in Palaeoanthropology.\u201d <em>British Journal for the History of Science<\/em> 42 (1): 95\u2013114.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Granger, Darryl E., Ryan J. Gibbon, Kathleen Kuman, Ronald J. Clarke, Laurent Bruxelles, and Marc W. Caffee. 2015. \u201cNew Cosmogenic Burial Ages for Sterkfontein Member 2 <em>Australopithecus<\/em> and Member 5 Oldowan.\u201d <em>Nature<\/em> 522 (7554): 85\u201388.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Granger Historical Picture Archive. 2018. <em>Cro-Magnon Footprint<\/em>. Image no. 0167868. Accessed March, 02, 2023. <a class=\"rId129\" href=\"https:\/\/www.granger.com\/results.asp?image=0167868&amp;itemw=4&amp;itemf=0001&amp;itemstep=1&amp;itemx=11&amp;screenwidth=1085\">https:\/\/www.granger.com\/results.asp?image=0167868&amp;itemw=4&amp;itemf=0001&amp;itemstep=1&amp;itemx=11&amp;screenwidth=1085<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Gruhn, R., 2020. \u201cEvidence Grows That Peopling of the Americas Began More Than 20,000 Years Ago.\u201d <em>Nature<\/em> 584: 47\u201348.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Haddy, A., and A. Hanson. 1982. \u201cResearch Notes and Application Reports Nitrogen and Fluorine Dating of Moundville Skeletal Samples. <em>Archaeometry<\/em> 24 (1): 37\u201344.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Hajdas, I., P. Ascough, M. H. Garnett, S. J. Fallon, C. L. Pearson, G. Quarta, K. L. Spalding, H. Yamaguchi, and M. Yoneda. 2021. \u201cRadiocarbon Dating.\u201d <em>Nature Reviews Methods Primers<\/em> 1 (62): 1\u201326.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Hatala, Kevin G., Brigitte Demes, and Brian G. Richmond. 2016. \u201cLaetoli Footprints Reveal Bipedal Gait Biomechanics Different from Those of Modern Humans and Chimpanzees.\u201d <em>Proceedings of the Royal Society B<\/em> 283: 20160235. <a class=\"rId130\" href=\"https:\/\/dx.doi.org\/10.1098\/rspb.2016.0235\">https:\/\/dx.doi.org\/10.1098\/rspb.2016.0235<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Hester, Thomas R., Harry J. Shafer, and Kenneth L. Feder. 1997. <em>Field Methods in Archaeology<\/em>, 7th ed. Mountain View, CA: Mayfield Publishing.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Hillam, J., C. M. Groves, D. M. Brown, M. G. L. Baillie, J. M. Coles, and B. J. Coles. 1990. \u201cDendrochronology of the English Neolithic.\u201d <em>Antiquity<\/em> 64 (243): 210\u2013220.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Holmes, Richard. 2010. <em>The Age of Wonder: How the Romantic Generation Discovered the Beauty and Terror of Science<\/em>. New York: Vintage.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Ingber, Sasha. 2012. \u201cVolcano Eruption Baked Rare Rhino Fossil.\u201d <em>National Geographic News<\/em>, November 30, 2012. Accessed July 25, 2018. https:\/\/www.nationalgeographic.com\/science\/article\/121130-rare-rhino-fossil-created-by-volcanic-explosion.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Jay, Mandy, B. T. Fuller, Michael P. Richards, Christopher J. Kn\u00fcsel, and Sarah S. King. 2008. \u201cIron Age Breastfeeding Practices in Britain: Isotopic Evidence from Wetwang Slack, East Yorkshire.\u201d <em>American Journal of Physical Anthropology<\/em> 136 (3): 327\u2013337.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Jones, Nicola. 2020. \u201cCarbon Dating, the Archaeological Workhorse, Is Getting a Major Reboot.\u201d <em>Nature<\/em><em> News<\/em>, May 19, 2020. doi: <a class=\"rId131\" href=\"https:\/\/doi.org\/10.1038\/d41586-020-01499-y\">https:\/\/doi.org\/10.1038\/d41586-020-01499-y<\/a><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Klein, Richard G. 1999. <em>The Human Career: Human Biological and Cultural Origins<\/em>, 2nd ed. Chicago: University of Chicago Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Komar, Debra A., and Jane E. Buikstra. 2008. <em>Forensic Anthropology: Contemporary Theory and Practice<\/em>. New York: Oxford University Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Kuniholm, Peter Ian, and Cecil L. Striker. 1987. \u201cDendrochronological Investigations in the Aegean and Neighboring Regions, 1983\u20131986.\u201d <em>Journal of Field Archaeology<\/em> 14 (4): 385\u2013398.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Laurenzi, Marinella A., Maria Laura Balestrieri, Giulio Bigazzi, Julio C. Hadler Neto, Pedro J. Iunes, Pio Norelli, Massimo Oddone, Ana Maria Osorio Araya, and Jos\u00e9 G. Viramonte. 2007. \u201cNew Constraints on Ages of Glasses Proposed as Reference Materials for Fission-Track Dating.\u201d <em>Geostandards &amp; Geoanalytical Research<\/em> 31 (2): 105\u2013124.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Lebatard, Anne-Elisabeth, Didier L. Bourl\u00e8s, Philippe Duringer, Marc Jolivet, R\u00e9gis Braucher, Julien Carcaillet, Mathieu Schuster, et al. 2008. \u201cCosmogenic Nuclide Dating of <em>Sahelanthropus tchadensis<\/em> and <em>Australopithecus bahrelghazali<\/em>: Mio-Pliocene Hominids from Chad.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 105 (9): 3226\u20133231.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Lyell, Charles. 1830\u20131833. <em>Principles of Geology, Being an Attempt to Explain the Former Changes of the Earth\u2019s Surface, by Reference to Causes Now in Operation<\/em>. 3 vols. London: John Murray.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Maixner, Frank, Dmitrij Turaev, Amaury Cazenave-Gassiot, Marek Janko, Ben Krause-Kyora, Michael R. Hoopmann, Ulrike Kusebauch, et al. 2018. \u201cThe Iceman\u2019s Last Meal Consisted of Fat, Wild Meat, and Cereals.\u201d <em>Current Biology<\/em> Report 28 (14): 2348\u20132355.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Masao, Fidelis T., Elgidius B. Ichumbaki, Marco Cherin, Angelo Barili, Giovanni Boschian, David A. Lurino, Sofia Menconero, Jacopo Moggi-Cecchi, and Giorgio Manzi. 2016. \u201cNew Footprints from Laetoli (Tanzania) Provide Evidence for Marked Body Size Variation in Early Hominins.\u201d\u00a0<em>eLife <\/em>5: e19568. <a class=\"rId132\" href=\"https:\/\/elifesciences.org\/articles\/19568\">https:\/\/elifesciences.org\/articles\/19568<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">McDonough, Katelyn, Taryn Johnson, Ted Goebel, and Karl Reinhard. 2022. \u201cDisease, Diet, and Thorny Headed Worm Infection in the Great Basin.\u201d Paper presented at 50th Annual Meeting of the Nevada Archaeological Association, Tonopah, Nevada, April 22nd, 2022.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Michels, Joseph W. 1972. \u201cDating Methods.\u201d <em>Annual Review of Anthropolog<\/em>y 1: 113\u2013126.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Monastersky, Richard. 2015. \u201cAnthropocene: The Human Age.\u201d <em>Nature<\/em> 519 (7542): 144\u2013147.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Montgomery, Janet, Jane A. Evans, Dominic Powlesland, and Charlotte A. Roberts. 2005. \u201cContinuity or Colonization in Anglo-Saxon England? Isotope Evidence for Mobility, Subsistence Practice, and Status at West Heslerton.\u201d <em>American Journal of Physical Anthropology<\/em> 126 (2): 123\u2013138.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">National Park Service. 2022. \u201cFossilized Footprints.\u201d <em>National Park Service <\/em>website, February 1. Accessed May 31, 2022. https:\/\/www.nps.gov\/whsa\/learn\/nature\/fossilized-footprints.htm.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Pastoors, Andreas, Tilman Lenssen-Erz, Bernd Breuckmann, Tsamkxao Ciqae, Ui Kxunta, Dirk Rieke-Zapp, and Thui Thao. 2017. \u201cExperience Based Reading of Pleistocene Human Footprints in Pech-Merle.\u201d <em>Quaternary International <\/em>430 (A): 155\u2013162.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Potts, Richard. 2012. \u201cEvolution and Environmental Change in Early Human Prehistory.\u201d <em>Annual Review of Anthropology<\/em> 41: 151\u2013167.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Raichlen, David A., and Adam D. Gordon. 2017. \u201cInterpretation of Footprints From Site S Confirms Human-like Bipedal Biomechanics in Laetoli Hominins.\u201d <em>Journal of Human Evolution<\/em> 107: 134\u2013138.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Ravn, Morten. 2010. \u201cBronze and Early Iron Age Bog Bodies from Denmark.\u201d <em>Acta Archaeologica<\/em> 81 (1): 112\u2013113.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Reimer, Paula J., Edouard Bard, Alex Bayliss, J. Warren Beck, Paul G. Blackwell, Christopher Bronk Ramsey, Caitlin E. Buck, et al. 2013. \u201cINTCAL13 and Marine13 Radiocarbon Age Calibration Curves 0\u201350,000 Years cal BP.\u201d <em>Radiocarbon<\/em> 55 (4): 1869\u20131887.<\/p>\n<p class=\"import-Normal\">Reinhard, Johan. 2006. <em>Ice Maiden: Inca Mummies, Mountain Gods, and Sacred Sites in the Andes<\/em>. Washington, DC.: National Geographic.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Reitz, Elizabeth J., and Elizabeth S. Wing. 1999. <em>Zooarchaeology<\/em>. Cambridge, UK: Cambridge University Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Renfrew, Colin, and Paul Bahn. 2016. <em>Archaeology: Theories, Methods, and Practice<\/em>, 7th ed. New York: Thames and Hudson.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Ruddiman, William F., Zhengtang Guo, Xin Zhou, Hanbin Wu, and Yanyan Yu. 2008. \u201cEarly Rice Farming and Anomalous Methane Trends.\u201d <em>Quaternary Science Reviews<\/em> 27 (13\u201314): 1291\u20131295.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Schmidt, Gavin. 1999. \u201cScience Briefs: Cold Climates, Warm Climates: How Can We Tell Past Temperatures?\u201d <em>National Aeronautics and Space Administration Goddard Institute for Space Studies<\/em>. <a class=\"rId133\" href=\"https:\/\/www.giss.nasa.gov\/research\/briefs\/1999_schmidt_01\/\">https:\/\/www.giss.nasa.gov\/research\/briefs\/1999_schmidt_01\/<\/a><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Shillito, Lisa-Marie, John C. Blong, Eleanor J. Green, and Eline N. van Asperen. 2020a. \u201cThe What, How and Why of Archaeological Coprolite Analysis.\u201d <em>Earth-Science Reviews<\/em> 207. <a class=\"rId134\" href=\"https:\/\/doi.org\/10.1016\/j.earscirev.2020.103196\">https:\/\/doi.org\/10.1016\/j.earscirev.2020.103196<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Shillito, Lisa-Marie, Helen L. Whelton, John C. Blong, Dennis L. Jenkins, Thomas J. Connolly, and Ian D. Bull. 2020b. \u201cPre-Clovis occupation of the Americas Identified by Human Fecal Biomarkers in Coprolites from Paisley Caves, Oregon.\u201d <em>Science Advances<\/em> 6 (29).\u200b\u200b https:\/\/www.science.org\/doi\/10.1126\/sciadv.aba6404.<\/p>\n<p class=\"import-Normal\">Sims, Douglas., and W. Spaulding. 2017. \u201cShallow Subsurface Evidence for Postglacial Holocene Lakes at Ivanpah Dry Lake: An Alternative Energy Development Site in the Central Mojave Desert, California, USA.\u201d <em>Journal of Geography and Geology<\/em> 9 (1): 1\u201324. DOI: <a class=\"rId135\" href=\"https:\/\/doi.org\/10.5539\/jgg.v9n1p1\">10.5539\/jgg.v9n1p1<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Slon, Viviane, Fabrizio Mafessoni, Benjamin Vernot, Cesare De Filippo, Steffi Grote, Bence Viola, Mateja Hajdinjak, St\u00e9phane Peyr\u00e9gne, Sarah Nagel, Samantha Brown, et al. 2018. \u201cThe Genome of the Offspring of a Neanderthal Mother and a Denisovan Father.\u201d <em>Nature<\/em> 561 (7721): 113\u2013116.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Smithsonian National Museum of Natural History. 2018. Laetoli Footprint Trails. Smithsonian National Museum of History, October 23. Accessed February 14, 2023. <a class=\"rId136\" href=\"https:\/\/humanorigins.si.edu\/evidence\/behavior\/footprints\/laetoli-footprint-trails\">https:\/\/humanorigins.si.edu\/evidence\/behavior\/footprints\/laetoli-footprint-trails<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Snoeck, Christophe, John Pouncett, Philippe Claeys, Steven Goderis, Nadine Mattielli, Mike Parker Pearson, Christie Willis, Antoine Zazzo, Julia A. Lee-Thorp, and Rick J. Schulting. 2018. \u201cStrontium Isotope Analysis on Cremated Human Remains from Stonehenge Support[sic] Links With West Wales.\u201d <em>Scientific Reports <\/em>8. <a class=\"rId137\" href=\"https:\/\/www.nature.com\/articles\/s41598-018-28969-8\">https:\/\/www.nature.com\/articles\/s41598-018-28969-8<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Spaulding, W. G., and D. B. Sims. 2018. \u201cA Glance to the East: Lake Ivanpah\u2014An Isolated Southern Great Basin Paleolake. <em>The 2018 Desert Symposium Field Guide and Proceedings<\/em>, 121-131. <a class=\"rId138\" href=\"https:\/\/www.researchgate.net\/publication\/340493333_A_glance_to_the_east_Lake_Ivanpah-_an_isolated_southern_Great_Basin_paleolake\">https:\/\/www.researchgate.net\/publication\/340493333_A_glance_to_the_east_Lake_Ivanpah-_an_isolated_southern_Great_Basin_paleolake<\/a><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Spray, Aaron. \u201cLa Brea Woman: Only (&amp; Controversial) Human from La Brea Tar Pits.\u201d <em>The Travel<\/em>, April 3, 2022. Accessed July 31, 2022. <a class=\"rId139\" href=\"https:\/\/www.thetravel.com\/what-to-know-of-the-la-brea-woman\/\">https:\/\/www.thetravel.com\/what-to-know-of-the-la-brea-woman\/<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Steffen, Will, Johan Rockstr\u00f6m, Katherine Richardson, Timothy M. Lenton, Carl Folke, Diana Liverman, Colin P. Summerhayes, et al. 2018. \u201cTrajectories of the Earth System in the Anthropocene.\u201d <em>PNAS <\/em>115 (33): 8252\u20138259.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Stodder, Ann L. W. 2008. \u201cTaphonomy and the Nature of Archaeological Assemblages.\u201d In <em>Biological Anthropology of the Human Skeleton<\/em>, edited by M. Anne Katzenberg and Shelley R. Saunders, 71\u2013115. Hoboken, NJ: Wiley Blackwell.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Straus, Lawrence Guy. 1989. \u201cGrave Reservations: More on Paleolithic Burial Evidence.\u201d <em>Current Anthropology<\/em> 30 (5): 633\u2013634.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Sullivan, Terry, and Harry Gifford. 1908. <em>She Sells Sea-Shells<\/em>. London: Francis, Day, and Hunter.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Taylor, Anthony, Jarod M. Hutson, Vaughn M. Bryant, and Dennis L. Jenkins. 2020. \u201cDietary Items in Early to Late Holocene Human Coprolites from Paisley Caves, Oregon, USA.\u201d <em>Palynology<\/em> 44 (1): 12\u201323.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Taylor, Paul D., Aaron O\u2019Dea. 2014. <em>A History of Life in 100 Fossils<\/em>. Washington, D.C.: Smithsonian Books.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">T\u00f6rnqvist, T. E., B. E. Rosenheim, P. Hu, and A. B. Fernandez. 2015. \u201cRadiocarbon Dating and Calibration.\u201d In <em>Handbook of Sea-Level Research<\/em>, edited by Ian Shennan, Antony J. Long, and Benjamin P. Horton, 347-360. Oxford, UK: John Wiley &amp; Sons. <a class=\"rId140\" href=\"https:\/\/doi.org\/10.1002\/9781118452547.ch23\">https:\/\/doi.org\/10.1002\/9781118452547.ch23<\/a><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">University of Arizona. n.d. \u201cUranium-Thorium Dating: The Uranium 238 Decay Series.\u201d Accessed November 21, 2022. <a class=\"rId141\" href=\"https:\/\/www.geo.arizona.edu\/Antevs\/ecol438\/uthdating.html\">https:\/\/www.geo.arizona.edu\/Antevs\/ecol438\/uthdating.html<\/a><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">University of the Witwatersrand. 2017. \u201cLittle Foot Takes a Bow: South Africa\u2019s Oldest and the World\u2019s Most Complete <em>Australopithecus<\/em> Skeleton Ever Found, Introduced to the World.\u201d <em>ScienceDaily<\/em>, December 6. Accessed February 14, 2023. <a class=\"rId142\" href=\"https:\/\/www.sciencedaily.com\/releases\/2017\/12\/171206100104.htm\">https:\/\/www.sciencedaily.com\/releases\/2017\/12\/171206100104.htm<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">van Calsteren, Peter, and Louise Thomas. 2006. \u201cUranium-Series Dating Applications in Natural Environmental Science.\u201d <em>Earth-Science Reviews<\/em> 75 (1\u20134): 155\u2013175.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Vanzetti, A., M. Vidale, M. Gallinaro, D. W. Frayer, and L. Bondioli. 2010. \u201cThe Iceman as a Burial.\u201d <em>Antiquity<\/em> 84 (325): 681\u2013692. <a class=\"rId143\" href=\"https:\/\/doi.org\/10.1017\/S0003598X0010016X\">https:\/\/doi.org\/10.1017\/S0003598X0010016X<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Verghese, Namrata. 2021. \u201cWhat Is Necropolitics? The Political Calculation of Life and Death.\u201d <em>Teen Vogue<\/em>. March 10, 2021. Accessed February 14, 2023. https:\/\/www.teenvogue.com\/story\/what-is-necropolitics.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Vidale, M., L. Bondioli, D. W. Frayer, M. Gallinaro, and A. Vanzetti. 2016. \u201c\u00d6tzi the Iceman.\u201d <em>Expedition<\/em> 58 (2): 13\u201317. Accessed February 14, 2023. <a class=\"rId144\" href=\"https:\/\/www.penn.museum\/sites\/expedition\/otzi-the-iceman\/\">https:\/\/www.penn.museum\/sites\/expedition\/otzi-the-iceman\/<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Wade, Lizzie. 2021. \"Footprints Support Claim of Early Arrival in the Americas.\" <em>Science <\/em>373 (6562): 1426. Accessed February 14, 2023. https:\/\/www.sciencemagazinedigital.org\/sciencemagazine\/24_september_2021\/MobilePagedArticle.action?articleId=1727132#articleId1727132.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Waters, Colin N., Jan Zalasiewicz, Anthony D. Barnosky, Alejandro Cearreta, Agieszka Galuszka, Juliana A. Ivar Do Sul, Catherine Jeandel, et al. 2016 \u201cIs the Anthropocene Distinct from the Holocene?\u201d <em>Science <\/em>351 (6269): aad2622-1-10. DOI:<a class=\"rId145\" href=\"https:\/\/dx.doi.org\/10.1126\/science.aad2622\">10.1126\/science.aad2622<\/a><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Watson, Traci. 2017. \u201cAncient Bones Reveal Girl\u2019s Tough Life in Early Americas.\u201d <em>Nature <\/em>544 (7648): 15\u201316<em>. <\/em><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Wendt, Kathleen, A., Xianglei Li,, and R. Lawrence Edwards. 2021. \u201cUranium-Thorium Dating of Speleothems.\u201d Elements 17 (2): 87\u201392.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">White, Tim D. 1986. \u201cCut Marks on the Bodo Cranium: A Case of Prehistoric Defleshing.\u201d <em>American Journal of Physical Anthropology<\/em> 69 (4): 503\u2013509.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Williams, Linda D. 2004. <em>Earth Science Demystified<\/em>. New York: McGraw-Hill Professional.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Wilson, Andrew S., Timothy Taylor, Maria Constanza Ceruti, Jose Antonio Chavez, Johan Reinhard, Vaughan Grimes, Wolfram Meier-Augenstein, et al. 2007. \u201cStable Isotope and DNA Evidence for Ritual Sequences in Inca Child Sacrifice.\u201d <em>PNAS<\/em> 104 (42): 16456\u201316461.<\/p>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">Acknowledgments<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\">We are grateful to Lee Anne Zajicek, who coauthored the first edition. Her original contributions continue to be an integral part of this chapter. We thank the staff of the Maturango Museum, Ridgecrest, California. Specifically, for their generous help with photography and fossil images, we acknowledge Debbie Benson, executive director; Alexander K. Rogers, former archaeology curator; Sherry Brubaker, natural history curator; and Elaine Wiley, history curator. We thank Sharlene Paxton, a librarian at Cerro Coso Community College, Ridgecrest, California, for her guidance and expertise with OER and open-source images, and John Stenger-Smith and Claudia Sellers from Cerro Coso Community College, Ridgecrest, California, for their feedback on the chemistry and plant biology content. Finally, we thank William Zajicek and Lauren Zajicek, our community college students, for providing their impressions and extensive feedback on early drafts of the chapter.<\/p>\n<\/div>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1806\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1806\"><div tabindex=\"-1\"><\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1772\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1772\"><div tabindex=\"-1\"><\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1773\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1773\"><div tabindex=\"-1\"><p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kerryn Warren, Ph.D., Grad Coach International<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lindsay Hunter, M.A., University of Iowa<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Navashni Naidoo, M.Sc., University of Cape Town<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Silindokuhle Mavuso, M.Sc., University of Witwatersrand<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\"><em>This chapter is a revision from <\/em><em>\"<\/em><a class=\"rId7\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\"><em>Chapter 9: Early Hominins<\/em><\/a><em>\" <\/em><em>by Kerryn Warren, K. Lindsay Hunter, Navashni Naidoo, Silindokuhle Mavuso, Kimberleigh Tommy, Rosa Moll, and Nomawethu Hlazo<\/em><em>. In <\/em><a class=\"rId8\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, first edition<\/em><\/a><em>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under <\/em><a class=\"rId9\" style=\"color: #000000\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\"><em>CC BY-NC 4.0<\/em><\/a><em>. <\/em><\/span><\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\"><span style=\"color: #000000\"><span style=\"color: #ffffff\">Learning Objectives<\/span><br \/>\n<\/span><\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li><span style=\"color: #000000\">Understand what is meant by \u201cderived\u201d and \u201cancestral\u201d traits and why this is relevant for understanding early hominin evolution.<\/span><\/li>\n<li><span style=\"color: #000000\">Understand changing paleoclimates and paleoenvironments as potential factors influencing early hominin adaptations.<\/span><\/li>\n<li><span style=\"color: #000000\">Describe the anatomical changes associated with bipedalism and dentition in early hominins, as well as their implications..<\/span><\/li>\n<li><span style=\"color: #000000\">Describe early hominin genera and species, including their currently understood dates and geographic expanses.<\/span><\/li>\n<li><span style=\"color: #000000\">Describe the earliest stone tool techno-complexes and their impact on the transition from early hominins to our genus.<\/span><\/li>\n<\/ul>\n<\/div>\n<\/div>\n<h2 class=\"__UNKNOWN__\"><span style=\"color: #000000\">Defining Hominins<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">It is through our study of our hominin ancestors and relatives that we are exposed to a world of \u201cmight have beens\u201d: of other paths not taken by our species, other ways of being human. But to better understand these different evolutionary trajectories, we must first define the terms we are using. If an imaginary line were drawn between ourselves and our closest relatives, the great apes, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1200\">bipedalism<\/a><\/strong> (or habitually walking upright on two feet) is where that line would be. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1270\">Hominin<\/a><\/strong>, then, means everyone on \u201cour\u201d side of the line: humans and all of our extinct bipedal ancestors and relatives since our divergence from the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1272\">last common ancestor (LCA)<\/a><\/strong> we share with chimpanzees.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Historic interpretations of our evolution, prior to our finding of early hominin <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1274\"><strong>fossils<\/strong><\/a>, varied. Debates in the mid-1800s regarding hominin origins focused on two key issues:<\/span><\/p>\n<ul>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">Where did we evolve?<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">Which traits evolved first?<\/span><\/li>\n<\/ul>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #00ff00\"><span style=\"background-color: #ccffcc\">Charles Darwin hypothesized that we evolved in Africa, as he was convinced that we shared greater commonality with chimpanzees and gorillas on the continent (Darwin 1871). Others, such as Ernst Haeckel and Eug\u00e8ne Dubois, insisted that we were closer in affinity to orangutans and that we evolved in Eurasia where, until the discovery of the Taung Child in South Africa in 1924, all humanlike fossils (of Neanderthals and <em>Homo erectus<\/em>) had been found (Shipman 2002).<\/span>\u00a0<span style=\"text-decoration: underline\">(<span style=\"background-color: #ff9900\">and refer to chapter)<\/span><\/span><\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Within this conversation, naturalists and early <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1276\">paleoanthropologists<\/a><\/strong> (people who study human evolution) speculated about which human traits came first. These included the evolution of a big brain (<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1278\"><strong>encephalization<\/strong><\/a>), the evolution of the way in which we move about on two legs (bipedalism), and the evolution of our flat faces and small teeth (indications of dietary change). Original hypotheses suggested that, in order to be motivated to change diet and move about in a bipedal fashion, the large brain needed to have evolved first, as is seen in the fossil species mentioned above.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">However, we now know that bipedal locomotion is one of the first things that evolved in our lineage, with early relatives having more apelike dentition and small brain sizes. While brain size expansion is seen primarily in our genus, <em>Homo<\/em>, earlier hominin brain sizes were highly variable between and within taxa, from 300 cc (cranial capacity, cm<sup>3<\/sup>), estimated in <em>Ardipithecus<\/em>, to 550 cc, estimated in <em>Paranthropus boisei<\/em>. The lower estimates are well within the range of variation of nonhuman extant great apes. In addition, body size variability also plays a role in the interpretation of whether brain size could be considered large or small for a particular species or specimen. In this chapter, we will tease out the details of early hominin evolution in terms of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1280\"><strong>morphology<\/strong><\/a> (i.e. the study of the form, size, or shape of things; in this case, skeletal parts).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">We also know that early human evolution occurred in a very complicated fashion. There were multiple species (multiple genera) that featured diversity in their diets and locomotion. Specimens have been found all along the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1282\"><strong>East African Rift System <\/strong>(<strong>EARS)<\/strong><\/a>; that is, in Ethiopia, Kenya, Tanzania, and Malawi; see Figure 9.1), in limestone caves in South Africa, and in Chad. Dates of these early relatives range from around 7 million years ago (mya) to around 1 mya, overlapping temporally with members of our genus, <em>Homo<\/em>.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 610px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/06\/image38.png\" alt=\"Patchy green mountain alongside a deep sandy valley in East Africa.\" width=\"610\" height=\"277\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.1: East African Rift System (EARS). Credit: <a href=\"https:\/\/www.flickr.com\/photos\/ninara\/8624605781\/in\/photolist-x2yH7-x2yHe-VfVWuD-e98mPF-SzzjsU-2bsBZhC-2hHec7m-xtJ7Ez-NXnXvh-7Yg3uo-2cS3FgG-2hjo1Dc-2hjGoTS-nnumi8-82U66W-dMNn7B-8jdVbd-NWDg8-NW6fj-ebhx5w-bkFv1G-Ct5ZD-5JQk8A-y6TgAc-x9k6oe-2ebLTDC-WcPMnJ-2ekh6CS-Cu3LH-xNHDFK-9RUsZi-94jVt4-P46uiB-QFyjyE-crU8N7-5JLJKV-2ekSgk8-5JL454-2cPgZrF-2bHfQZu-dMTVPN-6yUbeN-jzMicQ-48XjU9-2etR2Ze-Styrvw-crU7V7-2wakq3-crU6Z1-2etR2XR\/\">IMG_1696 Great Rift Valley<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/ninara\/\">Ninara<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\">CC BY 2.0 License<\/a>.<\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Yet there is still so much to understand. Modern debates now look at the relatedness of these species to us and to one another, and they consider which of these species were able to make and use tools. As a result, every <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1284\">site<\/a><\/strong> discovery in the patchy hominin fossil record tells us more about our evolution. In addition, recent scientific techniques (not available even ten years ago) provide new insights into the diets, environments, and lifestyles of these ancient relatives.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In the past, <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1288\"><strong>taxonom<\/strong><strong>y<\/strong><\/a> was primarily based on morphology. Today it is tied to known relationships based on molecular <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1290\"><strong>phylogeny<\/strong><\/a> (e.g., based on DNA) or a combination of the two. This is complicated when applied to living <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1286\"><strong>taxa<\/strong><\/a>, but becomes much more difficult when we try to categorize ancestor-descendant relationships for long-extinct species whose molecular information is no longer preserved. We therefore find ourselves falling back on morphological comparisons, often of teeth and partially fossilized skeletal material.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">It is here that we turn to the related concepts of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1292\">cladistics<\/a> <\/strong>and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1294\">phylogenetics<\/a><\/strong><strong>. <\/strong><span style=\"background-color: #ff99cc\">Cladistics groups organisms according to their last common ancestors based on shared <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1126\">derived traits<\/a><\/strong>. <\/span>In the case of early hominins, these are often morphological traits that differ from those seen in earlier populations. These new or modified traits provide evidence of evolutionary relationships, and organisms with the same derived traits are grouped in the same <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1116\">clade<\/a> <\/strong>(Figure 9.2). <span style=\"background-color: #ff99cc\">For example, if we use feathers as a trait, we can group pigeons and ostriches into the clade of birds.<\/span> In this chapter, we will examine the grouping of the Robust Australopithecines, whose cranial and dental features differ from those of earlier hominins, and therefore are considered derived.<\/span><\/p>\n<figure style=\"width: 708px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image48.png\" alt=\"Phylogenetic tree shows clades and non clade groupings.\" width=\"708\" height=\"192\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.2: Clades refer to groups of species or taxa that share a common ancestor. In <span class=\"ILfuVd\" lang=\"en\"><span class=\"hgKElc\">a phylogeny, a clade is a complete group of lineages, including their last common ancestor. Groupings that do not include a common ancestor and <em>all<\/em> of its descendants are not clades. <\/span><\/span>Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\">Clades (Figure 9.2)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Katie Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Dig Deeper: Problems Defining Hominin Species<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000\">It is worth noting that species designations for early hominin specimens are often highly contested. This is due to the fragmentary nature of the fossil record, the large timescale (millions of years) with which paleoanthropologists need to work, and the difficulty in evaluating whether morphological differences and similarities are due to meaningful phylogenetic or biological differences or subtle differences\/variation in niche occupation or time. In other words, do morphological differences really indicate different species? How would classifying species in the paleoanthropological record compare with classifying living species today, for whom we can sequence genomes and observe lifestyles?<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">There are also broader philosophical differences among researchers when it comes to paleo-species designations. Some scientists, known as \u201c<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1296\"><strong>lumpers<\/strong>,<\/a>\u201d argue that large variability is expected among multiple populations in a given species over time. These researchers will therefore prefer to \u201clump\u201d specimens of subtle differences into single taxa. Others, known as \u201c<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1298\"><strong>splitters<\/strong><\/a>,\u201d argue that species variability can be measured and that even subtle differences can imply differences in niche occupation that are extreme enough to mirror modern species differences. In general, splitters would consider geographic differences among populations as meaning that a species is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1300\">polytypic<\/a><\/strong> <span style=\"background-color: #ff99cc\">(i.e., capable of interacting and breeding biologically but having morphological population differences)<\/span>. This is worth keeping in mind when learning about why species designations may be contested.<\/span><\/p>\n<figure style=\"width: 520px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image8.jpg\" alt=\"A graph shows a curved line depicting changes in morphology among two species over time.\" width=\"520\" height=\"292\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.3: This graph demonstrates the concept of a chronospecies, where one species (Species A) \u201cevolves\u201d into another (Species B). Credit: Chronospecies original to Explorations: An Open Invitation to Biological Anthropology, 2nd edition by Kerryn Warren is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\"><span style=\"color: #000000\">This further plays a role in evaluating ancestry. Debates over which species \u201cgave rise\u201d to which continue to this day. It is common to try to create \u201clineages\u201d of species to determine when one species evolved into another over time. We refer to these as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1302\"><strong>chronospecies<\/strong><\/a> (Figure 9.3). Constructed hominin phylogenetic trees are routinely variable, changing with new specimen discoveries, new techniques for evaluating and comparing species, and, some have argued, nationalist or biased interpretations of the record. More recently, some researchers have shifted away from \u201ctreelike\u201d models of ancestry toward more nuanced metaphors such as the \u201cbraided stream,\u201d where some levels of interbreeding among species and populations are seen as natural processes of evolution.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Finally, it is worth considering the process of fossil discovery and publication. Some fossils are easily diagnostic to a species level and allow for easy and accurate interpretation. Some, however, are more controversial. This could be because they do not easily preserve or are incomplete, making it difficult to compare and place within a specific species (e.g., a fossil of a patella or knee bone). Researchers often need to make several important claims when announcing or publishing a find: a secure date (if possible), clear association with other finds, and an adequate comparison among multiple species (both extant and fossil). Therefore, it is not uncommon that an important find was made years before it is scientifically published.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Paleoenvironment and Hominin Evolution<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There is no doubt that one of the major selective pressures in hominin evolution is the environment. <span style=\"background-color: #ff9900\"><span style=\"background-color: #ffff00\">Large-scale changes in global and regional climate, as well as alterations to the environment, are<\/span> <\/span><em>(<span style=\"text-decoration: underline\">partially or have a great part<\/span>)<\/em><span style=\"background-color: #ffff00\"> all linked to hominin diversification, dispersal, and extinction<\/span> (Maslin et al. 2014). Environmental reconstructions often use modern analogues. Let us take, for instance, the hippopotamus. It is an animal that thrives in environments that have abundant water to keep its skin cool and moist. If the environment for some reason becomes drier, it is expected that hippopotamus populations will reduce. If a drier environment becomes wetter, it is possible that hippopotamus populations may be attracted to the new environment and thrive. Such instances have occurred multiple times in the past, and the bones of some <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1308\">fauna<\/a><\/strong> (i.e., animals, like the hippopotamus) that are sensitive to these changes give us insights into these events.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Yet reconstructing a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1304\">paleoenvironment<\/a><\/strong> relies on a range of techniques, which vary depending on whether research interests focus on local changes or more global environmental changes\/reconstructions. For local environments (such as a single site or region), comparing the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1306\">faunal assemblages<\/a> <\/strong>(collections of fossils of animals found at a site) with animals found in certain modern environments allows us to determine if past environments mirror current ones in the region. Changes in the faunal assemblages, as well as when they occur and how they occur, tell us about past environmental changes. Other techniques are also useful in this regard. Chemical analyses, for instance, can reveal the diets of individual fauna, providing clues as to the relative wetness or dryness of their environment (e.g., nitrogen <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1310\"><strong>isotopes<\/strong><\/a>; Kingston and Harrison 2007).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Global climatic changes in the distant past, which fluctuated between being colder and drier and warmer and wetter on average, would have global implications for environmental change (Figure 9.4). These can be studied by comparing marine core and terrestrial soil data across multiple sites. These techniques are based on chemical analysis, such as examination of the nitrogen and oxygen isotopes in shells and sediments. Similarly, analyzing pollen grains shows which kinds of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1312\"><strong>flora<\/strong> <\/a> survived in an environment at a specific time period. There are multiple lines of evidence that allow us to visualize global climate trends over millions of years (although it should be noted that the direction and extent of these changes could differ by geographic region).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 649px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image12-1-1.png\" alt=\"Chart shows cyclical carbon dioxide levels from 800,000 years ago until today.\" width=\"649\" height=\"406\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.4: This graph, based on the comparison of atmospheric samples contained in ice cores and more recent direct measurements, illustrates how atmospheric CO\u2082 has fluctuated over time and increased sharply since the Industrial Revolution. The graph also shows that since 800,000ya (and before) atmospheric CO\u2082 has never exceeded 300 parts per million (ppm). In 1950 it was 310ppm. Today atmospheric CO\u2082 has spiked to over 410 ppm. Credit: <a href=\"https:\/\/climate.nasa.gov\/evidence\/\">CO\u2082 increase since the Industrial Revolution<\/a> by <a href=\"https:\/\/www.nasa.gov\/\">NASA<\/a> is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a> and is used within <a href=\"https:\/\/www.nasa.gov\/multimedia\/guidelines\/index.html\">NASA guidelines on re-use<\/a>. Original from Luthi, D., et al.. 2008; Etheridge, D.M., et al. 2010; Vostok ice core data\/J.R. Petit et al.; NOAA Mauna Loa CO<a href=\"https:\/\/climate.nasa.gov\/evidence\/\">\u2082<\/a> record..<\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Both local and global climatic\/environmental changes have been used to understand factors affecting our evolution (DeHeinzelin et al. 1999; Kingston 2007). Environmental change acts as an important factor regarding the onset of several important hominin traits seen in early hominins and discussed in this chapter. Namely, the environment has been interpreted as the following:<\/span><\/p>\n<ul>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">the driving force behind the evolution of bipedalism,<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">the reason for change and variation in early hominin diets, and<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">the diversification of multiple early hominin species.<\/span><\/li>\n<\/ul>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are numerous hypotheses regarding how climate has driven and continues to drive human evolution. Here, we will focus on just three popular hypotheses.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Savannah Hypothesis (or Aridity Hypothesis)<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The hypothesis:<\/strong> This popular theory suggests that the expansion of the savannah (or less densely forested, drier environments) forced early hominins from an <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1314\"><strong>arboreal<\/strong> <\/a> lifestyle (one living in trees) to a terrestrial one where bipedalism was a more efficient form of locomotion (Figure 9.5). It was first proposed by Darwin (1871) and supported by anthropologists like Raymond Dart (1925). However, this idea was supported by little fossil or paleoenvironmental evidence and was later refined as the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1318\">Aridity Hypothesis<\/a><\/strong>. This hypothesis states that the long-term <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1316\">aridification<\/a><\/strong> and, thereby, expansion of savannah biomes were drivers in diversification in early hominin evolution (deMenocal 2004; deMenocal and Bloemendal 1995). It advocates for periods of accelerated aridification leading to early hominin speciation events.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 647px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image46.png\" alt=\"Photograph showing a dry, open savannah environment.\" width=\"647\" height=\"486\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.5: The African savannah grew during early hominin evolution. This may have forced early hominins from an arboreal lifestyle to a terrestrial one, where bipedalism was a more efficient form of locomotion. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:African_savannah_@_Masai_Mara_(21308330314).jpg\">African savannah @ Masai Mara (21308330314)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/132394214@N04\">Leo Li<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\">CC BY 2.0 License<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The evidence:<\/strong> While early bipedal hominins are often associated with wetter, more closed environments (i.e., not the Savannah Hypothesis), both marine and terrestrial records seem to support general cooling, drying conditions, with isotopic records indicating an increase in grasslands (i.e., colder and wetter climatic conditions) between 8 mya and 6 mya across the African continent (Cerling et al. 2011). This can be contrasted with later climatic changes derived from aeolian dust records (sediments transported to the site of interest by wind), which demonstrate increases in seasonal rainfall between 3 mya and 2.6 mya, 1.8 mya and 1.6 mya, and 1.2 mya and 0.8 mya (deMenocal 2004; deMenocal and Bloemendal 1995).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Interpretation(s):<\/strong> Despite a relatively scarce early hominin record, it is clear that two important factors occur around the time period in which we see increasing aridity. The first factor is the diversification of taxa, where high morphological variation between specimens has led to the naming of multiple hominin genera and species. The second factor is the observation that the earliest hominin fossils appear to have traits associated with bipedalism and are dated to around the drying period (as based on isotopic records). Some have argued that it is more accurately a combination of bipedalism and arboreal locomotion, which will be discussed later. However, the local environments in which these early specimens are found (as based on the faunal assemblages) do not appear to have been dry.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Turnover Pulse Hypothesis<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The hypothesis:<\/strong> In 1985, paleontologist Elisabeth Vbra noticed that in periods of extreme and rapid climate change, <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1326\"><strong>ungulates<\/strong><\/a> (hoofed mammals of various kinds) that had generalized diets fared better than those with specialized diets (Vrba 1988, 1998). <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1322\"><strong>Specialist<\/strong><\/a> eaters <span style=\"background-color: #ff99cc\">(those who rely primarily on specific food types)<\/span> faced extinction at greater rates than their <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1320\">generalist<\/a> <\/strong><span style=\"background-color: #ff99cc\">(those who can eat more varied and variable diets) <\/span>counterparts because they were unable to adapt to new environments (Vrba 2000). Thus, periods with extreme climate change would be associated with high <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1324\"><strong>faunal turnover<\/strong><\/a>: that is, the extinction of many species and the speciation, diversification, and migration of many others to occupy various niches.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The evidence:<\/strong> The onset of the<strong> <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1400\">Quaternary Ice Age<\/a><\/strong>, between 2.5 mya and 3 mya, brought extreme global, cyclical <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1328\"><strong>interglacial<\/strong> <\/a> and <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1330\"><strong>glacial<\/strong><\/a> periods (warmer, wetter periods with less ice at the poles, and colder, drier periods with more ice near the poles). Faunal evidence from the Turkana basin in East Africa indicates multiple instances of faunal turnover and extinction events, in which global climatic change resulted in changes from closed\/forested to open\/grassier habitats at single sites (Behrensmeyer et al. 1997; Bobe and Behrensmeyer 2004). Similarly, work in the Cape Floristic Belt of South Africa shows that extreme changes in climate play a role in extinction and migration in ungulates. While this theory was originally developed for ungulates, its proponents have argued that it can be applied to hominins as well. However, the link between climate and speciation is only vaguely understood (Faith and Behrensmeyer 2013).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Interpretation(s):<\/strong> While the evidence of rapid faunal turnover among ungulates during this time period appears clear, there is still some debate around its usefulness as applied to the paleoanthropological record. Specialist hominin species do appear to exist for long periods of time during this time period, yet it is also true that <em>Homo<\/em>, a generalist genus with a varied and adaptable diet, ultimately survives the majority of these fluctuations, and the specialists appear to go extinct.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Variability Selection Hypothesis<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The hypothesis: <\/strong>This hypothesis was first articulated by paleoanthropologist Richard Potts (1998). It links the high amount of climatic variability over the last 7 million years to both behavioral and morphological changes. Unlike previous notions, this hypothesis states that hominin evolution does not respond to habitat-specific changes or to specific aridity or moisture trends. Instead, long-term environmental unpredictability over time and space influenced morphological and behavioral adaptations that would help hominins survive, regardless of environmental context (Potts 1998, 2013). The Variability Selection Hypothesis states that hominin groups would experience varying degrees of natural selection due to continually changing environments and potential group isolation. This would allow certain groups to develop genetic combinations that would increase their ability to survive in shifting environments. These populations would then have a genetic advantage over others that were forced into habitat-specific adaptations (Potts 2013).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>The evidence:<\/strong> The evidence for this theory is similar to that for the Turnover Pulse Hypothesis: large climatic variability and higher survivability of generalists versus specialists. However, this hypothesis accommodates for larger time-scales of extinction and survival events.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Interpretation(s):<\/strong> In this way, the Variability Selection Hypothesis allows for a more flexible interpretation of the evolution of bipedalism in hominins and a more fluid interpretation of the Turnover Pulse Hypothesis, where species turnover is meant to be more rapid. In some ways, this hypothesis accommodates both environmental data and our interpretations of an evolution toward greater variability among species and the survivability of generalists.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000;background-color: #ff99cc\"><strong>Paleoenvironment Summary<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff99cc\">Some hypotheses presented in this section pay specific attention to habitat (Savannah Hypothesis) while others point to large-scale climatic forces (Variability Selection Hypothesis). Some may be interpreted to describe the evolution of traits such as bipedalism (Savannah Hypothesis), and others generally explain the diversification of early hominins (Turnover Pulse and Variability Selection Hypotheses). While there is no consensus as to how the environment drove our evolution, it is clear that the environment shaped both habitat and resource availability in ways that would have influenced our early ancestors physically and behaviorally.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Derived Adaptations: Bipedalism<br \/>\n<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The unique form of locomotion exhibited by modern humans, called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1332\">obligate bipedalism<\/a><\/strong>, is important in distinguishing our species from the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1334\"><strong>extant<\/strong><\/a> (living) great apes. The ability to walk habitually upright is thus considered one of the defining attributes of the hominin lineage. We also differ from other animals that walk bipedally (such as kangaroos) in that we do not have a tail to balance us as we move.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The origin of bipedalism in hominins has been debated in paleoanthropology, but at present there are two main <span style=\"background-color: #ff99cc\">ideas<\/span>: <span style=\"text-decoration: underline\">(<em>theories<\/em>)<\/span><\/span><\/p>\n<ol>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">early hominins initially lived in trees, but increasingly started living on the ground, so we were a product of an arboreal last common ancestor (LCA) or,<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\">our LCA was a terrestrial quadrupedal knuckle-walking species, more similar to extant chimpanzees.<\/span><\/li>\n<\/ol>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Most research supports the first theory of an arboreal LCA based on skeletal morphology of early hominin genera that demonstrate adaptations for climbing but not for knuckle-walking. This would mean that both humans and chimpanzees can be considered \u201cderived\u201d in terms of locomotion since chimpanzees would have independently evolved knuckle-walking.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are many current ideas regarding selective pressures that would lead to early hominins adapting upright posture and locomotion. Many of these selective pressures, as we have seen in the previous section, coincide with a shift in environmental conditions, supported by paleoenvironmental data. In general, however, it appears that, like extant great apes, early hominins thrived in forested regions with dense tree coverage, which would indicate an arboreal lifestyle. As the environmental conditions changed and a savannah\/grassland environment became more widespread, the tree cover would become less dense, scattered, and sparse such that bipedalism would become more important.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are several proposed selective pressures for bipedalism:<\/span><\/p>\n<ol>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\"><strong>E<\/strong><strong>nergy conservation:<\/strong> Modern bipedal humans conserve more energy than extant chimpanzees, which are predominantly knuckle-walking quadrupeds when walking over land. While chimpanzees, for instance, are faster than humans terrestrially, they expend large amounts of energy being so. Adaptations to bipedalism include \u201cstacking\u201d the majority of the weight of the body over a small area around the center of gravity (i.e., the head is above the chest, which is above the pelvis, which is over the knees, which are above the feet). This reduces the amount of muscle needed to be engaged during locomotion to \u201cpull us up\u201d and allows us to travel longer distances expending far less energy.<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\"><strong>T<\/strong><strong>hermoregulation:<\/strong> Less surface area (i.e., only the head and shoulders) is exposed to direct sunlight during the hottest parts of the day (i.e., midday). This means that the body has less need to employ additional \u201ccooling\u201d mechanisms such as sweating, which additionally means less water loss.<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent;text-align: left;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Bipedalism <\/strong><span style=\"text-decoration: underline\">(Freeing of Hands)<\/span><strong>: <\/strong>This method of locomotion freed up our ancestors\u2019 hands such that they could more easily gather food and carry tools or infants. This further enabled the use of hands for more specialized adaptations associated with the manufacturing and use of tools.<\/span><\/li>\n<\/ol>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">These selective pressures are not mutually exclusive. Bipedality could have evolved from a combination of these selective pressures, in ways that increased the chances of early hominin survival.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Skeletal Adaptations for Bipedalism<\/strong><\/span><\/h3>\n<figure style=\"width: 405px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image10-1.png\" alt=\"A full human skeleton and gorilla skeleton standing in upright positions next to each other.\" width=\"405\" height=\"452\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 9.6: Compared to gorillas (right) and other apes, humans (left) have highly specialized adaptations to facilitate bipedal locomotion. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Primatenskelett-drawing.jpg\">Skeleton of human (1) and gorilla (2), unnaturally sketched<\/a> by unknown from Brehms Tierleben, Small Edition 1927 is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<br \/><\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Humans have highly specialized adaptations to facilitate obligate bipedalism (Figure 9.6). Many of these adaptations occur within the soft tissue of the body (e.g., muscles and tendons). However, when analyzing the paleoanthropological record for evidence of the emergence of bipedalism, all that remains is the fossilized bone. Interpretations of locomotion are therefore often based on comparative analyses between fossil remains and the skeletons of extant primates with known locomotor behaviors. These adaptations occur throughout the skeleton and are summarized in Figure 9.7.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The majority of these adaptations occur in the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1336\"><strong>postcranium<\/strong><\/a> <span style=\"background-color: #ff99cc\">(the skeleton from below the head)<\/span> and are outlined in Figure 9.7. In general, these adaptations allow for greater stability and strength in the lower limb, by allowing for more shock absorption, for a larger surface area for muscle attachment, and for the \u201cstacking\u201d of the skeleton directly over the center of gravity to reduce energy needed to be kept upright. These adaptations often mean less flexibility in areas such as the knee and foot.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">However, these adaptations come at a cost. Evolving from a nonobligate bipedal ancestor means that the adaptations we have are evolutionary compromises. For instance, the valgus knee (angle at the knee) is an essential adaptation to balance the body weight above the ankle during bipedal locomotion. However, the strain and shock absorption at an angled knee eventually takes its toll. For example, runners often experience joint pain. Similarly, the long neck of the femur absorbs stress and accommodates for a larger pelvis, but it is a weak point, resulting in hip replacements being commonplace among the elderly, especially in cases where the bone additionally weakens through osteoporosis. Finally, the S-shaped curve in our spine allows us to stand upright, relative to the more curved C-shaped spine of an LCA. Yet the weaknesses in the curves can lead to pinching of nerves and back pain. Since many of these problems primarily are only seen in old age, they can potentially be seen as an evolutionary compromise.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Despite relatively few postcranial fragments, the fossil record in early hominins indicates a complex pattern of emergence of bipedalism. Key features, such as a more anteriorly placed foramen magnum, are argued to be seen even in the earliest discovered hominins, indicating an upright posture (Dart 1925). Some early species appear to have a mix of ancestral (arboreal) and derived (bipedal) traits, which indicates a mixed locomotion and a more <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1402\">mosaic evolution<\/a><\/strong> of the trait. Some early hominins appear to, for instance, have bowl-shaped pelvises (hip bones) and angled femurs suitable for bipedalism but also have retained an opposable <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1338\">hallux<\/a><\/strong> (big toe) or curved fingers and longer arms (for arboreal locomotion). These mixed morphologies may indicate that earlier hominins were not fully obligate bipeds and thus thrived in mosaic environments.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Yet the associations between postcranial and the more diagnostic cranial fossils and bones are not always clear, muddying our understanding of the specific species to which fossils belong (Grine et al. 2022).<\/span><\/p>\n<div align=\"left\">\n<table class=\"grid\">\n<caption>\n<p class=\"import-Normal\" style=\"text-align: left\"><span style=\"color: #000000\">Figure 9.7: Skeletal comparisons between modern humans (obligate bipeds) and nonobligate bipeds (e.g., chimpanzees). Credit: <a class=\"rId34\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\">Skeletal comparisons between modern humans and <\/a><a class=\"rId35\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\">nonobligate<\/a><a class=\"rId36\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\"> bipeds (Figure 9.6)<\/a> original to <a class=\"rId37\" style=\"color: #000000\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a class=\"rId38\" style=\"color: #000000\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/span><\/p>\n<\/caption>\n<thead>\n<tr>\n<td style=\"width: 97.998px\"><strong>Region<\/strong><\/td>\n<td style=\"width: 106.992px\"><strong>Feature<\/strong><\/td>\n<td style=\"width: 366.992px\"><strong>Obligate Biped (H. sapiens)<\/strong><\/td>\n<td style=\"width: 310px\"><strong>Nonobligate Biped<\/strong><\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr>\n<td style=\"width: 97.998px\">Cranium<\/td>\n<td style=\"width: 106.992px\">Position of the foramen magnum<\/td>\n<td style=\"width: 366.992px\">Positioned inferiorly (immediately under the cranium) so that the head rests on top of the vertebral column for balance and support (head is perpendicular to the ground).<\/td>\n<td style=\"width: 310px\">Posteriorly positioned (to the back of the cranium). Head is positioned parallel to the ground.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Body proportions<\/td>\n<td style=\"width: 366.992px\">Shorter upper limb (not used for locomotion).<\/td>\n<td style=\"width: 310px\">Longer upper limbs (used for locomotion).<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Spinal curvature<\/td>\n<td style=\"width: 366.992px\">S-curve due to pressure exerted on the spine from bipedalism (lumbar lordosis).<\/td>\n<td style=\"width: 310px\">C-curve.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Vertebrae<\/td>\n<td style=\"width: 366.992px\">Robust lumbar (lower-back) vertebrae (for shock absorbance and weight bearing). Lower back is more flexible than that of apes as the hips and trunk swivel when walking (weight transmission).<\/td>\n<td style=\"width: 310px\">Gracile lumbar vertebrae compared to those of modern humans.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Pelvis<\/td>\n<td style=\"width: 366.992px\">Shorter, broader, bowl-shaped pelvis (for support); very robust. Broad sacrum with large sacroiliac joint surfaces.<\/td>\n<td style=\"width: 310px\">Longer, flatter, elongated ilia; more narrow and gracile; narrower sacrum; relatively smaller sacroiliac joint surface.<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Lower limb<\/td>\n<td style=\"width: 366.992px\">In general, longer, more robust lower limbs and more stable, larger joints.<\/p>\n<ul>\n<li style=\"font-weight: 400\">Large femoral head and longer neck (absorbs more stress and increases the mechanical advantage).<\/li>\n<li style=\"font-weight: 400\">Valgus knee, in which the angle of the knee positions it over the ankle and keeps the center of gravity balanced over the stance leg during stride cycle (shock absorbance).<\/li>\n<li style=\"font-weight: 400\">Distal tibia (lower leg) of humans has a large medial malleolus for stability.<\/li>\n<\/ul>\n<\/td>\n<td style=\"width: 310px\">In general, smaller, more gracile limbs with more flexible joints.<\/p>\n<ul>\n<li style=\"font-weight: 400\">Femoral neck is smaller in comparison to modern humans and shorter.<\/li>\n<li style=\"font-weight: 400\">The legs bow outward, and there is no valgus angle of the knee (no \u201cknock knees\u201d).<\/li>\n<li style=\"font-weight: 400\">The distal tibia in chimpanzees is trapezoid (wider anteriorly) for climbing and allows more flexibility.<\/li>\n<\/ul>\n<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 97.998px\">Post<\/p>\n<p>cranium<\/td>\n<td style=\"width: 106.992px\">Foot<\/td>\n<td style=\"width: 366.992px\">Rigid, robust foot, without a midtarsal break.<\/p>\n<p>Nonopposable and large, robust big toe (for push off while walking) and large heel for shock absorbance.<\/td>\n<td style=\"width: 310px\">Flexible foot, midtarsal break present (which allows primates to lift their heels independently from their feet), opposable big toe for grasping.<\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<p class=\"import-Normal\"><span style=\"color: #000000;background-color: #ccffcc\">It is also worth noting that, while not directly related to bipedalism per se, other postcranial adaptations are evident in the hominin fossil record from some of the earlier hominins. For instance, the hand and finger morphologies of many of the earliest hominins indicate adaptations consistent with arboreality. These include longer hands, more curved metacarpals and phalanges (long bones in the hand and fingers, respectively), and a shorter, relatively weaker thumb. This allows for gripping onto curved surfaces during locomotion. The earliest hominins appear to have mixed morphologies for both bipedalism and arborealism. However, among Australopiths (members of the genus, Australopithecus), there are indications for greater reliance on bipedalism as the primary form of locomotion. Similarly, adaptations consistent with tool manufacture (shorter fingers and a longer, more robust thumb, in contrast to the features associated with arborealism) have been argued to appear before the genus <em>Homo<\/em>.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Early Hominins: Sahelanthropus and Orrorin<\/strong><\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">We see evidence for bipedalism in some of the earliest fossil hominins, dated from within our estimates of our divergence from chimpanzees. These hominins, however, also indicate evidence for arboreal locomotion.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The earliest dated hominin find (between 6 mya and 7 mya, based on radiometric dating of volcanic tufts) has been argued to come from Chad and is named <strong><em>Sahelanthropus tchadensis<\/em> <\/strong>(Figure 9.8; Brunet et al. 1995). The initial discovery was made in 2001 by Ahounta Djimdoumalbaye and announced in <em>Nature<\/em> in 2002 by a team led by French paleontologist Michel Brunet. The find has a small cranial capacity (360 cc) and smaller canines than those in extant great apes, though they are larger and pointier than those in humans. <span style=\"background-color: #ffff00\">This implies strongly that, over evolutionary time, the need for display and dominance among males has reduced, as has our sexual dimorphism<\/span>. A short cranial base and a foramen magnum that is more humanlike in positioning have been argued to indicate upright walking.<\/span><\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 640px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-289\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.82.jpg\" alt=\"Four views of a beige-colored skull are seen against a black background.\" width=\"640\" height=\"640\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.8: Sahelanthropus tchadensis exhibits a set of derived features, including a long, low cranium; a small, ape-sized braincase; and relatively reduced prognathism. Credit: aa <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 anterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 posterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 inferior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; and d <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Sahelanthropus%20tchadensis\/TM%20266-01-060-1\">Sahelanthropus tchadensis: TM 266-01-060-1 lateral left view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Initially, the inclusion of <em>Sahelanthropus<\/em> in the hominin family was debated by researchers, since the evidence for bipedalism is based on cranial evidence alone, which is not as convincing as postcranial evidence. Yet, a femur (thigh bone) and ulnae (upper arm bones) thought to belong to <em>Sahelanthropus<\/em> was discovered in 2001 (although not published until 2022). These bones may support the idea that the hominin was in fact a terrestrial biped with arboreal capabilities and behaviors (Daver et al. 2022).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Orrorin tugenensis<\/em><\/strong> (Orrorin meaning \u201coriginal man\u201d), dated to between 6 mya and 5.7 mya, was discovered near Tugen Hills in Kenya in 2000. Smaller <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1340\">cheek teeth<\/a><\/strong> (molars and premolars) than those in even more recent hominins, thick enamel, and reduced, but apelike, canines characterize this species. This is the first species that clearly indicates adaptations for bipedal locomotion, with fragmentary leg, arm, and finger bones having been found but few cranial remains. One of the most important elements discovered was a proximal femur, BAR 1002'00. The femur is the thigh bone, and the proximal part is that which articulates with the pelvis; this is very important for studying posture and locomotion. This femur indicates that <em>Ororrin<\/em> was bipedal, and recent studies suggest that it walked in a similar way to later <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1404\">Pliocene<\/a><\/strong> hominins. Some have argued that features of the finger bones suggest potential tool-making capabilities, although many researchers argue that these features are also consistent with climbing.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Early Hominins: The Genus <em>Ardipithecus<\/em><\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Another genus, <em>Ardipithecus<\/em>, is argued to be represented by at least two species: <em>Ardipithecus (Ar.) ramidus <\/em>and <em>Ar. kadabba<\/em>.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Ardipithecus ramidus<\/em><\/strong> (\u201cramid\u201d means root in the Afar language) is currently the best-known of the earliest hominins (Figure 9.9). Unlike <em>Sahelanthropus<\/em> and<em> Orrorin<\/em>, this species has a large sample size of over 110 specimens from Aramis alone. Dated to 4.4 mya, <em>Ar. ramidus<\/em> was found in Ethiopia (in the Middle Awash region and in Gona). This species was announced in 1994 by American palaeoanthropologist Tim White, based on a partial female skeleton nicknamed \u201cArdi\u201d (ARA-VP-6\/500; White et al. 1994). Ardi demonstrates a mosaic of ancestral and derived characteristics in the postcrania. For instance, she had an opposable big toe (hallux), similar to chimpanzees (i.e., more ancestral), which could have aided in climbing trees effectively. However, the pelvis and hip show that she could walk upright (i.e., it is derived), supporting her hominin status. A small brain (300 cc to 350 cc), midfacial projection, and slight prognathism show retained ancestral cranial features, but the cheek bones are less flared and robust than in later hominins.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 706px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-1310\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.9-scaled-1.jpg\" alt=\"Skull cast and partial skeleton with photographs of some bones and line drawings of others.\" width=\"706\" height=\"453\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.9a and b: Researchers believe that Ardipithecus ramidus was able to walk upright, although not as efficiently as later humans. It possessed the musculature required for tree climbing, and while moving quadrupedally, it likely placed weight on the palms of the hands rather than on the knuckles. Credit: a. <a class=\"rId61\" href=\"https:\/\/boneclones.com\/product\/ardipithecus-ramidus-skull-BH-039\">Ardipithecus ramidus Skull<\/a> by <a class=\"rId62\" href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a> is used by permission and available here under a <a class=\"rId63\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>; b. <a class=\"rId64\" href=\"https:\/\/boneclones.com\/product\/ardipithecus-ramidus-skull-BH-039\">Artist\u2019s rendition of \u201cArdi\u201d skeleton<\/a> by <a class=\"rId65\" href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a> is used by permission and available here under a <a class=\"rId66\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong><em>Ardipithecus kadabba<\/em><\/strong> (the species name means \u201coldest ancestor\u201d in the Afar language) is known from localities on the western margin of the Middle Awash region, the same locality where <em>Ar. ramidus<\/em> has been found. Specimens include mandibular fragments and isolated teeth as well as a few postcranial elements from the Asa Koma (5.5 mya to 5.77 mya) and Kuseralee Members (5.2 mya), well-dated and understood (but temporally separate) volcanic layers in East Africa. This species was discovered in 1997 by paleoanthropologist Dr. Yohannes Haile-Selassie. Originally these specimens were referred to as a subspecies of <em>Ar. ramidus<\/em>. In 2002, six teeth were discovered at Asa Koma and the dental-wear patterns confirmed that this was a distinct species, named <em>Ar. kadabba,<\/em> in 2004. One of the postcranial remains recovered included a 5.2 million-year-old toe bone that demonstrated features that are associated with toeing off (pushing off the ground with the big toe leaving last) during walking, a characteristic unique to bipedal walkers. However, the toe bone was found in the Kuseralee Member, and therefore some doubt has been cast by researchers about its association with the teeth from the Asa Koma Member.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"background-color: #ff99cc\"><strong><span style=\"color: #000000\">Bipedal Trends in Early Hominins: Summary<\/span><\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff99cc\">Trends toward bipedalism are seen in our earliest hominin finds. However, many specimens also indicate retained capabilities for climbing. Trends include a larger, more robust hallux; a more compact foot, with an arch; a robust, long femur, angled at the knee; a robust tibia; a bowl-shaped pelvis; and a more anterior foramen magnum. While the level of bipedality in <em>Salehanthropus<\/em> <em>tchadenisis<\/em> is debated since there are few fossils and no postcranial evidence, <em>Orrorin tugenensis<\/em> and <em>Ardipithecus<\/em> <em>k<\/em><em>adabba <\/em>show clear indications of some of these bipedal trends. However, some retained ancestral traits, such as an opposable hallux in <em>Ardipithecus<\/em>, indicate some retention in climbing ability.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Derived Adaptations: Early Hominin Dention<\/span><\/h2>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">The Importance of Teeth<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Teeth are abundant in the fossil record, primarily because they are already highly mineralized as they are forming, far more so than even bone. Because of this, teeth preserve readily. And, because they preserve readily, they are well-studied and better understood than many skeletal elements. In the sparse hominin (and primate) fossil record, teeth are, in some cases, all we have.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Teeth also reveal a lot about the individual from whom they came. We can tell what they evolved to eat, to which other species they may be closely related, and even, to some extent, the level of sexual dimorphism, or general variability, within a given species. This is powerful information that can be contained in a single tooth. With a little more observation, the wearing patterns on a tooth can tell us about the diet of the individual in the weeks leading up to its death. Furthermore, the way in which a tooth is formed, and the timing of formation, can reveal information about changes in diet (or even mobility) over infancy and childhood, using isotopic analyses. When it comes to our earliest hominin relatives, this information is vital for understanding how they lived.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The purpose of comparing different hominin species is to better understand the functional morphology as it applies to dentition. In this, we mean that the morphology of the teeth or masticatory system (which includes jaws) can reveal something about the way in which they were used and, therefore, the kinds of foods these hominins ate. When comparing the features of hominin groups, it is worth considering modern analogues (i.e., animals with which to compare) to make more appropriate assumptions about diet. In this way, hominin dentition is often compared with that of chimpanzees and gorillas (our close ape relatives), as well as with that of modern humans.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The most divergent group, however, is humans. Humans around the world have incredibly varied diets. Among hunter-gatherers, it can vary from a honey- and plant-rich diet, as seen in the Hadza in Tanzania, to a diet almost entirely reliant on animal fat and protein, as seen in Inuits in polar regions of the world. We are therefore considered generalists, more general than the largely <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1342\"><strong>frugivorous<\/strong><\/a> (fruit-eating) chimpanzee or the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1344\"><strong>folivorous<\/strong><\/a> (foliage-eating) gorilla, as discussed in Chapter 5.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">One way in which all humans are similar is our reliance on the processing of our food. We cut up and tear meat with tools using our hands, instead of using our front teeth (incisors and canines). We smash and grind up hard seeds, instead of crushing them with our hind teeth (molars). This means that, unlike our ape relatives, we can rely more on developing tools to navigate our complex and varied diets. <span style=\"text-decoration: underline\">(We could say)<\/span> Our brain, therefore, is our primary masticatory organ. Evolutionarily, our teeth have reduced in size and our faces are flatter, or more <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1346\">orthognathic<\/a>, <\/strong>partially in response to our increased reliance on our hands and brain to process food. Similarly, a reduction in teeth and a more generalist dental morphology could also indicate an increase in softer and more variable foods, such as the inclusion of more meat. <span style=\"background-color: #ffff00\">These trends begin early on in our evolution.<\/span> The link has been made between some of the earliest evidence for stone tool manufacture, the earliest members of our genus, and the features that we associate with these specimens.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">General Dental Trends in Early Hominins<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Several trends are visible in the dentition of early hominins. However, all tend to have the same <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1348\"><strong>dental formula<\/strong><\/a>. The dental formula tells us how many of each tooth type are present in each quadrant of the mouth. Going from the front of the mouth, this includes the square, flat <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1160\">incisors<\/a><\/strong>; the pointy <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1166\">canines<\/a><\/strong>; the small, flatter <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1350\"><strong>premolars<\/strong><\/a>; and the larger hind <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1164\">molars<\/a><\/strong>. In many primates, from Old World monkeys to great apes, the typical dental formula is 2:1:2:3. This means that if we divide the mouth into quadrants, each has two incisors, one canine, two premolars, and three molars. The eight teeth per quadrant total 32 teeth in all (although some humans have fewer teeth due to the absence of their wisdom teeth, or third molars).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 380px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image24.png\" alt=\"Anterior view of the lower face of a person showing their teeth.\" width=\"380\" height=\"253\" \/><figcaption class=\"wp-caption-text\">Figure 9.10: In humans, our canines are often a similar size to our incisors. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Adult_human_teeth.jpg\">Adult human teeth<\/a> by <a href=\"https:\/\/www.genusfotografen.se\/\">Genusfotografen<\/a> (Tomas Gunnarsson) through <a href=\"https:\/\/wikimedia.se\/\">Wikimedia Sverige<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p><span style=\"color: #000000\">The morphology of the individual teeth is where we see the most change. Among primates, large incisors are associated with food procurement or preparation (such as biting small fruits), while small incisors indicate a diet that may contain small seeds or leaves (where the preparation is primarily in the back of the mouth). Most hominins have relatively large, flat, vertically aligned incisors that <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1396\">occlude<\/a> <\/strong>(touch) relatively well, forming a \u201cbite.\u201d This differs from, for instance, the orangutan, whose teeth stick out (i.e.<em>,<\/em> are <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1352\"><strong>procumbent<\/strong><\/a>).<\/span><\/p>\n<p>&nbsp;<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">While the teeth are often aligned with diet, the canines may be misleading in that regard. We tend to associate pointy, large canines with the ripping required for meat, and the reduction (or, in some animals, the absence) of canines as indicative of herbivorous diets. In humans, our canines are often a similar size to our incisors and therefore considered <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1398\">incisiform<\/a><\/strong> (Figure 9.10). However, our closest relatives all have very long, pointy canines, particularly on their upper dentition. This is true even for the gorilla, which lives almost exclusively on plants. The canines in these instances reveal more about social structure and sexual dimorphism than diet, as large canines often signal <span style=\"background-color: #ffff00\">dominance<\/span>.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Early on in human evolution, we see a reduction in canine size. <em>Sahelanthropus tchadensis<\/em> and <em>Orrorin tugenensis<\/em> both have smaller canines than those in extant great apes, yet the canines are still larger and pointier than those in humans or more recent hominins. <span style=\"background-color: #ffff00\">This implies strongly that, over evolutionary time, the need for display and dominance among males has reduced, as has our sexual dimorphism.<\/span> In <em>Ardipithecus ramidus<\/em>, there is no obvious difference between male and female canine size, yet they are still slightly larger and pointier than in modern humans. This implies a less sexually dimorphic social structure in the earlier hominins relative to modern-day chimpanzees and gorillas.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Along with a reduction in canine size is the reduction or elimination of a canine <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1172\">diastema<\/a>:<\/strong> a gap between the teeth on the mandible that allows room for elongated teeth on the maxilla to \u201cfit\u201d in the mouth. Absence of a diastema is an excellent indication of a reduction in canine size. In animals with large canines (such as baboons), there is also often a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1354\"><strong>honing P3<\/strong><\/a>, where the first premolar (also known as P3 for evolutionary reasons) is triangular in shape, \u201csharpened\u201d by the extended canine from the upper dentition. This is also seen in some early hominins: <em>Ardipithecus<\/em>, for example, has small canines that are almost the same height as its incisors, although still larger than those in recent hominins.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The hind dentition, such as the bicuspid (two cusped) premolars or the much larger molars, are also highly indicative of a generalist diet in hominins. Among the earliest hominins, the molars are larger than we see in our genus, increasing in size to the back of the mouth and angled in such a way from the much smaller anterior dentition as to give these hominins a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1356\"><strong>parabolic<\/strong><\/a> (V-shaped) dental arch. This differs from our living relatives and some early hominins, such as <em>Sahelanthropus<\/em>, whose molars and premolars are relatively parallel between the left and right sides of the mouth, creating a U-shape.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Among more recent early hominins, the molars are larger than those in the earliest hominins and far larger than those in our own genus, <em>Homo.<\/em> Large, short molars with thick <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1358\"><strong>enamel<\/strong><\/a> allowed our early cousins to grind fibrous, coarse foods, such as sedges, which require plenty of chewing. This is further evidenced in the low <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1176\">cusps<\/a>,<\/strong> or ridges, on the teeth, which are ideal for chewing. In our genus, the hind dentition is far smaller than in these early hominins. Our teeth also have medium-size cusps, which allow for both efficient grinding and tearing\/shearing meats.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Understanding the dental morphology has allowed researchers to extrapolate very specific behaviors of early hominins. It is worth noting that while teeth preserve well and are abundant, a slew of other morphological traits additionally provide evidence for many of these hypotheses. Yet there are some traits that are ambiguous. For instance, while there are definitely high levels of sexual dimorphism in <em>Au. afarensis<\/em>, discussed in the next section, the canine teeth are reduced in size, implying that while canines may be useful indicators for sexual dimorphism, it is also worth considering other evidence.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000;background-color: #ff99cc\">In summary, trends among early hominins include a reduction in procumbency, reduced hind dentition (molars and premolars), a reduction in canine size (more incisiform with a lack of canine diastema and honing P3), flatter molar cusps, and thicker dental enamel. All early hominins have the ancestral dental formula of 2:1:2:3. These trends are all consistent with a generalist diet, incorporating more fibrous foods.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Special Topic: Contested Species<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Many named species are highly debated and argued to have specimens associated with a more variable <em>Au. afarensis <\/em>or <em>Au. anamensis<\/em> species. Sometimes these specimens are dated to times when, or found in places in which, there are \u201cgaps\u201d in the palaeoanthropological record. These are argued to represent chronospecies or variants of <em>Au. afarensis<\/em>. However, it is possible that, with more discoveries, the distinct species types will hold.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><strong><em>Australopithecus bahrelghazali<\/em><\/strong> is dated to within the time period of <em>Au. afarensi<\/em>s (3.6 mya; Brunet et al. 1995) and was the first Australopithecine to be discovered in Chad in central Africa. Researchers argue that the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1360\">holotype<\/a><\/strong>, whom discoverers have named \u201cAbel,\u201d falls under the range of variation of <em>Au. afarensis<\/em> and therefore that <em>A. bahrelghazali<\/em> does not fall into a new species (Lebatard et al. 2008). If \u201cAbel\u201d is a member of <em>Au. afarensis<\/em>, the geographic range of the species would be greatly extended.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">On a different note, <strong><em>Australopithecus <\/em><\/strong><strong><em>deyiremada<\/em><\/strong> (meaning \u201cclose relative\u201d in the Ethiopian language of Afar) is dated to 3.5 mya to 3.3 mya and is based on fossil mandible bones discovered in 2011 in Woranso-Mille (in the Afar region of Ethiopia) by Yohannes Haile-Selassie, an Ethiopian paleoanthropologist (Haile-Selassie et al. 2019). The discovery indicated, in contrast to <em>Au. afarensis<\/em>, smaller teeth with thicker enamel (potentially suggesting a harder diet) as well as a larger mandible and more projecting cheekbones. This find may be evidence that more than one closely related hominin species occupied the same region at the same temporal period (Haile-Selassie et al. 2015; Spoor 2015) or that other <em>Au. afarensis<\/em> specimens have been incorrectly designated. However, others have argued that this species has been prematurely identified and that more evidence is needed before splitting the taxa, since the variation appears subtle and may be due to slightly different niche occupations between populations over time.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><strong><em>Australopithecus garhi<\/em><\/strong> is another species found in the Middle Awash region of Ethiopia. It is currently dated to 2.5 mya (younger than <em>Au. afarensis<\/em>). Researchers have suggested it fills in a much-needed temporal \u201cgap\u201d between hominin finds in the region, with some anatomical differences, such as a relatively large cranial capacity (450 cc) and larger hind dentition than seen in other gracile Australopithecines. Similarly, the species has been argued to have longer hind limbs than <em>Au. afarensis<\/em>, although it was still able to move arboreally (Asfaw et al. 1999). However, this species is not well documented or understood and is based on only several fossil specimens. More astonishingly, crude stone tools resembling Oldowan (which will be described later) have been found in association with <em>Au. garhi<\/em>. While lacking some of the features of the Oldowan, this is one of the earliest technologies found in direct association with a hominin.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><strong><em>Kenyanthopus<\/em><\/strong><strong><em> platyops<\/em><\/strong> (the name \u201cplatyops\u201d refers to its flatter-faced appearance) is a highly contested genus\/species designation of a specimen (KNM-WT 40000) from Lake Turkana in Kenya, discovered by Maeve Leakey in 1999 (Figure 9.11). Dated to between 3.5 mya and 3.2 mya, some have suggested this specimen is an <em>Australopithecus<\/em>, perhaps even <em>Au.<\/em> <em>afarensis<\/em> (with a brain size which is difficult to determine, yet appears small), while still others have placed this specimen in <em>Homo <\/em>(small dentition and flat-orthognathic face). While taxonomic placing of this species is quite divided, the discoverers have argued that this species is ancestral to <em>Homo<\/em>, in particular to <em>Homo <\/em><em>ruldolfensis<\/em> (Leakey et al. 2001). Some researchers have additionally associated the earliest tool finds from Lomekwi, Kenya, temporally (3.3 mya) and in close geographic proximity to this specimen.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 579px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-292 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.11.jpg\" alt=\"Four views of an ancient skull are shown on a black background.\" width=\"579\" height=\"579\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.11: This specimen, KNM WT 40000 (Kenyanthopus platyops), has small detention, a small brain case, and a relatively flat face. Its genus\/species designation remains contested. Credit: a. <a class=\"rId76\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId77\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 anterior view<\/a> by \u00a9<a class=\"rId78\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId79\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId80\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId81\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId82\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 superior view<\/a> by \u00a9<a class=\"rId83\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId84\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId85\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a class=\"rId86\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId87\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 lateral left view<\/a> by \u00a9<a class=\"rId88\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId89\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId90\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; d. <a class=\"rId91\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"><em>Kenyanthropus platyops<\/em><\/a><a class=\"rId92\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Kenyanthropus%20platyops\/KNM%20WT%2040000\"> KNM WT 40000 inferior view<\/a> by \u00a9<a class=\"rId93\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId94\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId95\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">The Genus <em>Australopithecus<\/em><br \/>\n<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The Australopithecines are a diverse group of hominins, comprising various species. <em>Australopithecus<\/em> is the given group or genus name. It stems from the Latin word <em>Australo<\/em>, meaning \u201csouthern,\u201d and the Greek word <em>pithecus,<\/em> meaning \u201cape.\u201d Within this section, we will outline these differing species\u2019 geological and temporal distributions across Africa, unique derived and\/or shared traits, and importance in the fossil record.<\/span><\/p>\n<figure style=\"width: 381px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image37-2.png\" alt=\"A skull has a pronounced sagittal crest, flaring cheekbones, and large hind teeth.\" width=\"381\" height=\"585\" \/><figcaption class=\"wp-caption-text\">Figure 9.12: Robust Australopithecines such as Paranthropus boisei had large molars and chewing muscles. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Paranthropus_boisei_skull.jpg\">Paranthropus boisei skull<\/a> by Durova is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/deed.en\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Between 3 mya and 1 mya, there seems to be differences in dietary strategy between different species of hominins designated as Australopithecines. A pattern of larger posterior dentition (even relative to the incisors and canines in the front of the mouth), thick enamel, and cranial evidence for extremely large chewing muscles is far more pronounced in a group known as the robust australopithecines. This pattern is extreme<span style=\"text-decoration: underline\">ly<\/span> relative to their earlier contemporaries or predecessors, the gracile australopithecines<strong>,<\/strong> and is certainly larger than those seen in early <em>Homo<\/em>, which emerged during this time. This pattern of incredibly large hind dentition (and very small anterior dentition) has led people to refer to robust australopithecines as <strong>megadont<\/strong> hominins (Figure 9.12).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Because of these differences, this section has been divided into \u201cgracile\u201d and \u201crobust\u201d Australopithecines, highlighting the morphological differences between the two groups (which many researchers have designated as separate genera: <em>Australopithecus<\/em> and <em>Paranthropus<\/em>, respectively) and then focusing on the individual species. It is worth noting, however, that not all researchers accept these clades as biologically or genetically distinct, with some researchers insisting that the relative gracile and robust features found in these species are due to parallel evolutionary events toward similar dietary niches.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Despite this genus\u2019 ancestral traits and small cranial capacity, all members show evidence of bipedal locomotion. It is generally accepted that <em>Australopithecus <\/em>species display varying degrees of arborealism along with bipedality.<\/span><\/p>\n<h3 class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Gracile Australopithecines<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">This section describes individual species from across Africa. These species are called \u201c<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1406\">gracile<\/a> <\/strong>australopithecines\u201d because of their smaller and less robust features compared to the divergent \u201c<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1408\"><strong>robust<\/strong><\/a>\u201d group. Numerous Australopithecine species have been named, but some are only based on a handful of fossil finds, whose designations are controversial.<\/span><\/p>\n<h4 class=\"import-Normal\"><em><span style=\"color: #000000\">East African Australopithecines<\/span><\/em><\/h4>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">East African Australopithecines are found throughout the EARS, and they include the earliest species associated with this genus. Numerous fossil-yielding sites, such as Olduvai, Turkana, and Laetoli, have excellent, datable stratigraphy, owing to the layers of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1362\"><strong>volcanic tufts<\/strong> <\/a> that have accumulated over millions of years. <span style=\"background-color: #ff9900\">These tufts may be dated using absolute dating techniques, such as Potassium-Argon dating (described in Chapter 7). This means that it is possible to know a relatively refined date for any fossil if the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1364\"><strong>context<\/strong> <\/a> (i.e., exact location) of that find is known.<\/span> Similarly, comparisons between the faunal assemblages of these stratigraphic layers have allowed researchers to chronologically identify environmental changes.<\/span><\/p>\n<figure style=\"width: 313px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image20-1-1.png\" alt=\"Occlusal view of an Au. anamensis mandible, with relatively large teeth, including canines.\" width=\"313\" height=\"313\" \/><figcaption class=\"wp-caption-text\">Figure 9.13: As seen in this mandible of KNM-KP 29281, Australopithecus anamensis had relatively large canine teeth. Credit: <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20anamensis\/KNM-KP%2029281\">Australopithecus anamensis: KNM-KP 29281 occlusal view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p><span style=\"color: #000000\">The earliest known Australopithecine is dated to 4.2 mya to 3.8 mya. <strong><em>Australopithecus anamensis<\/em><\/strong> (after \u201cAnam,\u201d meaning \u201clake\u201d from the Turkana region in Kenya; Leakey et al. 1995; Patterson and Howells 1967) is currently found from sites in the Turkana region (Kenya) and Middle Awash (Ethiopia; Figure 9.13). Recently, a 2019 find from Ethiopia, named MRD, after Miro Dora where it was found, was discovered by an Ethiopian herder named Ali Bereino. It is one of the most complete cranial finds of this species (Ward et al. 1999). A small brain size (370 cc), relatively large canines, projecting cheekbones, and earholes show more ancestral features as compared to those of more recent Australopithecines. The most important element discovered with this species is a fragment of a tibia (shinbone), which demonstrates features associated with weight transfer during bipedal walking. Similarly, the earliest found hominin femur belongs to this species. Ancestral traits in the upper limb (such as the humerus) indicate some retained arboreal locomotion.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Some researchers suggest that <em>Au. anamensis<\/em> is an intermediate form of the chronospecies that becomes <em>Au. afarensis<\/em>, evolving from <em>Ar. ramidus<\/em>. However, this is debated, with other researchers suggesting morphological similarities and affinities with more recent species instead. Almost 100 specimens, representing over 20 individuals, have been found to date (Leakey et al. 1995; McHenry 2009; Ward et al. 1999).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Au. afarensis<\/em><\/strong> is one of the oldest and most well-known australopithecine species and consists of a large number of fossil remains. <em>Au. afarensis<\/em> (which means \u201cfrom the Afar region\u201d) is dated to between 2.9 mya and 3.9 mya and is found in sites all along the EARS system, in Tanzania, Kenya, and Ethiopia (Figure 9.14). The most famous individual from this species is a partial female skeleton discovered in Hadar (Ethiopia), later nicknamed \u201cLucy,\u201d after the Beatles\u2019 song \u201cLucy in the Sky with Diamonds,\u201d which was played in celebration of the find (Johanson et al. 1978; Kimbel and Delezene 2009). This skeleton was found in 1974 by Donald Johanson and dates to approximately 3.2 mya. In addition, in 2002 a juvenile of the species was found by Zeresenay Alemseged and given the name \u201cSelam\u201d (meaning \u201cpeace,\u201d DIK 1-1), though it is popularly known as \u201cLucy\u2019s Child\u201d or as the \u201cDikika Child\u201d (Alemseged et al. 2006). Similarly, the \u201cLaetoli Footprints\u201d (discussed in Chapter 7; Hay and Leakey 1982; Leakey and Hay 1979) have drawn much attention.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 643px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-295 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.14.jpg\" alt=\"Two images of life-like reconstructions of female and male Au. afarensis.\" width=\"643\" height=\"322\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.14 a-b: Artistic reconstructions of Australopithecus afarensis by artist John Gurche. Female \u201cLucy\u201d is left and a male is on the right. Credit: a. <a class=\"rId106\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Australopithecus afarensis, \u201cLucy,\u201d adult female. Reconstruction based on AL-288-1 by artist John Gurche, front view close-up<\/a> by <a class=\"rId107\" href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?\u201d] is <a class=\"rId108\" href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>; b. <a class=\"rId109\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Australopithecus afarensis, adult male. Reconstruction based on <\/a><a class=\"rId110\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">AL444-2<\/a><a class=\"rId111\" href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\"> by John Gurche<\/a> by <a class=\"rId112\" href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?\u201d] is <a class=\"rId113\" href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>.<\/figcaption><\/figure>\n<figure style=\"width: 320px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image40.png\" alt=\"A partial skeleton includes bones of the cranium, mandible, and postcranium.\" width=\"320\" height=\"772\" \/><figcaption class=\"wp-caption-text\">Figure 9.15: The humanlike femoral neck, valgus knee, and bowl-shaped hip seen in the \u201cLucy\u201d skeleton indicates that Australopithecus afarensis was bipedal. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Lucy_blackbg.jpg\">Lucy blackbg<\/a> [AL 288-1, Australopithecus afarensis, cast from Museum national d'histoire naturelle, Paris] by 120 is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The canines and molars of <em>Au. afarensis<\/em> are reduced relative to great apes but are larger than those found in modern humans (indicative of a generalist diet); in addition, <em>Au. afarensis <\/em>has a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1366\"><strong>prognathic<\/strong> <\/a> face (the face below the eyes juts anteriorly) and robust facial features that indicate relatively strong chewing musculature (compared with <em>Homo<\/em>) but which are less extreme than in <em>Paranthropus<\/em>. Despite a reduction in canine size in this species, large overall size variation indicates high levels of sexual dimorphism.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Skeletal evidence indicates that this species was bipedal, as its pelvis and lower limb demonstrate a humanlike femoral neck, valgus knee, and bowl-shaped hip (Figure 9.15). <span style=\"background-color: #ccffcc\">More evidence of bipedalism is found in the footprints of this species<\/span>. <em>Au. afarensis<\/em> is associated with the Laetoli Footprints, <span style=\"background-color: #ff9900\">a 24-meter trackway of hominin fossil footprints preserved in volcanic ash discovered by Mary Leakey in Tanzania and dated to 3.5 mya to 3 mya. This set of prints is thought to have been produced by three bipedal individuals as there are no knuckle imprints, no opposable big toes, and a clear arch is present. The infants of this species are thought to have been more arboreal than the adults, as discovered through analyses of the foot bones of the Dikika Child dated to 3.32 mya (Alemseged et al. 2006).<\/span><\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Although not found in direct association with stone tools, potential evidence for cut marks on bones, found at Dikika, and dated to 3.39 mya indicates a possible temporal\/ geographic overlap between meat eating, tool use, and this species. However, this evidence is fiercely debated. Others have associated the cut marks with the earliest tool finds from Lomekwi, Kenya, temporally (3.3 mya) and in close geographic proximity to this species.<\/span><\/p>\n<h4 class=\"import-Normal\"><em><span style=\"color: #000000\">South African Australopithecines<\/span><\/em><\/h4>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Since the discovery of the Taung Child, there have been numerous Australopithecine discoveries from the region known as \u201cThe Cradle of Humankind,\u201d which was recently given UNESCO World Heritage Site status as \u201cThe Fossil Hominid Sites of South Africa.\u201d The limestone caves found in the Cradle allow for the excellent preservation of fossils. Past animals navigating the landscape and falling into cave openings, or caves used as dens by carnivores, led to the accumulation of deposits over millions of years. Many of the hominin fossils, encased in <strong>breccia<\/strong> (hard, calcareous sedimentary rock), are recently exposed from limestone quarries mined in the previous century. This means that extracting fossils requires excellent and detailed exposed work, often by a team of skilled technicians.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">While these sites have historically been difficult to date, with mixed assemblages accumulated over large time periods, advances in techniques such as uranium-series dating have allowed for greater accuracy. Historically, the excellent faunal record from East Africa has been used to compare sites based on <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1368\"><strong>relative dating<\/strong>,<\/a> whereby environmental and faunal changes and extinction events allow us to know which hominin finds are relatively younger or older than others.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The discovery of the Taung Child in 1924 (discussed in the Special Topic box \u201cThe Taung Child\u201d below) shifted the focus of palaeoanthropological research from Europe to Africa, although acceptance of this shift was slow (Broom 1947; Dart 1925). The species to which it is assigned, <strong><em>Australopithecus africanus<\/em><\/strong> (name meaning \u201cSouthern Ape of Africa\u201d), is currently dated to between 3.3 mya and 2.1 mya (Pickering and Kramers 2010), with discoveries from Sterkfontein, Taung, Makapansgat, and Gladysvale in South Africa (Figure 9.16). A relatively large brain (400 cc to 500 cc), small canines without an associated diastema, and more rounded cranium and smaller teeth than <em>Au. afarensis<\/em> indicate some derived traits. Similarly, the postcranial remains (in particular, the pelvis) indicate bipedalism. However, the sloping face and curved phalanges (indicative of retained arboreal locomotor abilities) show some ancestral features. Although not in direct association with stone tools, a 2015 study noted that the trabecular bone morphology of the hand was consistent with forceful tool manufacture and use, suggesting potential early tool abilities.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 570px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image19-1.png\" alt=\"A life-like reconstruction of the face of Australopithecus africanus, smiling in anterior view.\" width=\"570\" height=\"570\" \/><figcaption class=\"wp-caption-text\">Figure 9.16: An artistic reconstruction of Australopithecus africanus by John Gurche. Credit: <a href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Australopithecus africanus. Reconstruction based on STS 5 by John Gurche <\/a>by <a href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?] is <a href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Another famous <em>Au. africanus<\/em> skull (the skull of \u201cMrs. Ples\u201d) was previously attributed to <em>Plesianthropus transvaalensis<\/em><em>, <\/em>meaning \u201cnear human from the Transvaal,\u201d the old name for Gauteng Province, South Africa (Broom 1947, 1950). The name was shortened by contemporary journalists to \u201cPles\u201d (Figure 9.17). Due to the prevailing mores of the time, the assumed female found herself married, at least in name, and has become widely known as \u201cMrs. Ples.\u201d It was later reassigned to <em>Au. africanus<\/em> and is now argued by some to be a young male rather than an adult female cranium (Thackeray 2000, Thackeray et al. 2002).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 548px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-298 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.17.jpg\" alt=\"Four views of an ancient skull are shown on a black background.\" width=\"548\" height=\"548\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.17: The \u201cMrs. Ples\u201d brain case is small in size (like apes) but its face is less prognathic; its foramen magnum is positioned more like a modern human than an African apes. Credit: a. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 anterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 posterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 superior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; and d. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Sts%205\">Australopithecus africanus Sts 5 lateral right view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In 2008, nine-year-old Matthew Berger, son of paleoanthropologist Lee Berger, noted a clavicle bone in some leftover mining breccia in the Malapa Fossil Site (South Africa). After rigorous studies, the species, <strong><em>Australopithecus sediba<\/em><\/strong> (meaning \u201cfountain\u201d or \u201cwellspring\u201d in the South African language of Sesotho), was named in 2010 (Figure 9.18; Berger et al. 2010). The first type specimen belongs to a juvenile male, Karabo (MH1), but the species is known from at least six partial skeletons, from infants through adults. These specimens are currently dated to 1.97 mya (Dirks et al. 2010). The discoverers have argued that <em>Au. sediba<\/em> shows mosaic features between <em>Au. africanus<\/em> and the genus, <em>Homo<\/em>, which potentially indicates a transitional species, although this is heavily debated. These features include a small brain size (<em>Australopithecus<\/em>-like; 420 cc to 450 cc) but gracile mandible and small teeth (<em>Homo<\/em>-like). Similarly, the postcranial skeletons are also said to have mosaic features: scientists have interpreted this mixture of traits (such as a robust ankle but evidence for an arch in the foot) as a transitional phase between a body previously adapted to arborealism (particularly in evidence from the bones of the wrist) to one that adapted to bipedal ground walking. Some researchers have argued that <em>Au. sediba<\/em> shows a modern hand morphology (shorter fingers and a longer thumb), indicating that adaptations to tool manufacture and use may be present in this species.<\/span><\/p>\n<figure style=\"width: 531px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image17-1.png\" alt=\"A beige-colored skull with no mandible on a black background has some missing teeth.\" width=\"531\" height=\"400\" \/><figcaption class=\"wp-caption-text\">Figure 9.18: Australopithecus sediba shows mosaic features between Au. africanus and Homo. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Australopithecus_sediba.JPG\">Australopithecus sediba<\/a>, photo by Brett Eloff courtesy <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Profberger\">Profberger<\/a> and <a href=\"https:\/\/en.wikipedia.org\/wiki\/University_of_the_Witwatersrand\">Wits University<\/a>, is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Another famous Australopithecine find from South Africa is that of the nearly complete skeleton now known as \u201cLittle Foot\u201d (Clarke 1998, 2013). Little Foot (StW 573) is potentially the earliest dated South African hominin fossil, dating to 3.7 mya, based on radiostopic techniques, although some argue that it is younger than 3 mya (Pickering and Kramers 2010). The name is jokingly in contrast to the cryptid species \u201cbigfoot\u201d and is named because the initial discovery of four ankle bones indicated bipedality. Little Foot was discovered by Ron Clarke in 1994, when he came across the ankle bones while sorting through monkey fossils in the University of Witwatersrand collections (Clarke and Tobias 1995). He asked Stephen Motsumi and Nkwane Molefe to identify the known records of the fossils, which allowed them to find the rest of the specimen within just days of searching the Sterkfontein Caves\u2019 Silberberg Grotto.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The discoverers of Little Foot insist that other fossil finds, previously identified as <em>Au. Africanus<\/em>, be placed in this new species based on shared ancestral traits with older East African Australopithecines (Clarke and Kuman 2019). These include features such as a relatively large brain size (408 cc), robust zygomatic arch, and a flatter midface. Furthermore, the discoverers have argued that the heavy anterior dental wear patterns, relatively large anterior dentition, and smaller hind dentition of this specimen more closely resemble that of <em>Au. anamensis<\/em> or <em>Au. afarensis<\/em>. It has thus been placed in the species <strong><em>Australopithecus prometheus<\/em><\/strong>. This species name refers to a previously defunct taxon named by Raymond Dart. The species designation was, through analyzing Little Foot, revived by Ron Clarke, who insists that many other fossil hominin specimens have prematurely been placed into <em>Au. africanus<\/em>. Others say that it is more likely that <em>Au. africanus<\/em> is a more variable species and not representative of two distinct species.<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\"><em>Paranthropus<\/em> \u201cRobust\u201d Australopithecines<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In the robust australopithecines, the specialized nature of the teeth and masticatory system, such as flaring zygomatic arches (cheekbones), accommodate very large temporalis (chewing) muscles. These features also include a large, broad, dish-shaped face and and a large mandible with extremely large posterior dentition (referred to as megadonts) and hyper-thick enamel (Kimbel 2015; Lee-Thorp 2011; Wood 2010). Research has revolved around the shared adaptations of these \u201crobust\u201d australopithecines, linking their morphologies to a diet of hard and\/or tough foods (Brain 1967; Rak 1988). Some argued that the diet of the robust australopithecines was so specific that any change in environment would have accelerated their extinction. The generalist nature of the teeth of the gracile australopithecines, and of early <em>Homo<\/em>, would have made them more capable of adapting to environmental change. However, some have suggested that the features of the robust australopithecines might have developed as an effective response to what are known as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1370\"><strong>fallback <\/strong><strong>foods<\/strong><\/a> in hard times rather than indicating a lack of adaptability.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">There are currently three widely accepted robust australopithecus or, <em>Paranthropus<\/em>, species: <em>P. aethiopic<\/em><em>us<\/em>, which has more ancestral traits, and <em>P. boisei and P. robustus<\/em>, which are more derived in their features (Strait et al. 1997; Wood and Schroer 2017). These three species have been grouped together by a majority of scholars as a single genus as they share more derived features (are more closely related to each other; or, in other words, are <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1372\"><strong>monophyletic<\/strong><\/a>) than the other australopithecines (Grine 1988; Hlazo 2015; Strait et al. 1997; Wood 2010 ). While researchers have mostly agreed to use the umbrella term <em>Paranthropus<\/em>, there are those who disagree (Constantino and Wood 2004, 2007; Wood 2010).<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">As a collective, this genus spans 2.7 mya to 1.0 mya, although the dates of the individual species differ. The earliest of the Paranthropus species, <strong><em>Paranthropus aethiopicus<\/em><\/strong>, is dated to between 2.7 mya and 2.3 mya and currently found in Tanzania, Kenya, and Ethiopia in the EARS system (Figure 9.19; Constantino and Wood 2007; Hlazo 2015; Kimbel 2015; Walker et al. 1986; White 1988). It is well known because of one specimen known as the \u201cBlack Skull\u201d (KNM\u2013WT 17000), so called because of the mineral manganese that stained it black during fossilization (Kimbel 2015). As with all robust Australopithecines, <em>P. aethiopicus<\/em> has the shared derived traits of large, flat premolars and molars; large, flaring zygomatic arches for accommodating large chewing muscles (the temporalis muscle); a sagittal crest (ridge on the top of the skull) for increased muscle attachment of the chewing muscles to the skull; and a robust mandible and supraorbital torus (brow ridge). However, only a few teeth have been found. A proximal tibia indicates bipedality and similar body size to <em>Au. afarensis<\/em>. In recent years, researchers have discovered and assigned a proximal tibia and juvenile cranium (L.338y-6) to the species (Wood and Boyle 2016).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 666px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-300 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.19.jpg\" alt=\"Five views of a beige partial skull on a black background.\" width=\"666\" height=\"444\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.19: The \u201cBlack Skull\u201d (Paranthropus aethiopicus) had a large sagittal crest and large, flared zygomatic arches that indicate it had large chewing muscles and a powerful biting force. Credit: a. <a class=\"rId156\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId157\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 anterior view<\/a> by \u00a9<a class=\"rId158\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId159\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId160\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId161\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId162\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 lateral right view<\/a> by \u00a9<a class=\"rId163\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId164\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId165\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a class=\"rId166\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId167\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 superior view<\/a> by \u00a9<a class=\"rId168\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId169\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId170\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; d. <a class=\"rId171\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId172\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 posterior view<\/a> by \u00a9<a class=\"rId173\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId174\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId175\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; e. <a class=\"rId176\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\"><em>Paranthropus aethiopicus<\/em><\/a><a class=\"rId177\" href=\"https:\/\/efossils.org\/page\/boneviewer\/paranthropus%20aethiopicus\/KNM-WT%2017000\">: KNM-WT 17000 inferior view<\/a> by \u00a9<a class=\"rId178\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId179\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId180\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">First attributed as <em>Zinjanthropus boisei<\/em> (with the first discovery going by the nickname \u201cZinj\u201d or sometimes \u201cNutcracker Man\u201d), <strong><em>Paranthropus boisei<\/em><\/strong> was discovered in 1959 by Mary Leakey (see Figure 9.20 and 9.21; Hay 1990; Leakey 1959). This \u201crobust\u201d australopith species is distributed across countries in East Africa at sites such as Kenya (Koobi Fora, West Turkana, and Chesowanja), Malawi (Malema-Chiwondo), Tanzania (Olduvai Gorge and Peninj), and Ethiopia (Omo River Basin and Konso). The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1374\">hypodigm<\/a><\/strong>, sample of fossils whose features define the group, has been found by researchers to date to roughly 2.4 mya to 1.4 mya. Due to the nature of its exaggerated, larger, and more robust features, <em>P. boisei <\/em>has been termed <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1376\"><strong>hyper-robust<\/strong><\/a>\u2014that is, even more heavily built than other robust species, with very large, flat posterior dentition (Kimbel 2015). Tools dated to 2.5 mya in Ethiopia have been argued to possibly belong to this species. Despite the cranial features of <em>P. boisei<\/em> indicating a tough diet of tubers, nuts, and seeds, isotopes indicate a diet high in C4 foods (e.g., grasses, such as sedges). Another famous specimen from this species is the Peninj mandible from Tanzania, found in 1964 by Kimoya Kimeu.<\/span><\/p>\n<figure style=\"width: 557px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image14-1.png\" alt=\"Life-like reconstruction of Paranthropus boisei.\" width=\"557\" height=\"557\" \/><figcaption class=\"wp-caption-text\">Figure 9.20: Artistic reconstruction of a Paranthropus boisei, male, by John Gurche. Credit: <a href=\"https:\/\/humanorigins.si.edu\/multimedia\/slideshows\/reconstructed-faces\">Paranthropus boisei, male. Reconstruction based on OH 5 and KNM-ER 406 by John Gurche<\/a> by <a href=\"https:\/\/www.si.edu\/\">the Smithsonian<\/a> [exhibit: \u201cReconstructed Faces: What Does It Mean to Be Human?\u201d] is <a href=\"https:\/\/www.si.edu\/termsofuse\/\">copyrighted and used for educational and noncommercial purposes as outlined by the Smithsonian<\/a>.<\/figcaption><\/figure>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 565px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-302 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.21.jpg\" alt=\"Three views of an ancient skull are shown on a black background.\" width=\"565\" height=\"565\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.21: \u201cNutcracker Man\u201d (Paranthropus boisei) had hyper-robust features including very large dentition, flaring zygomatic arches, a broad concave face. It had a powerful and extremely efficient chewing force. Credit: <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20boisei\/OH%205\">Paranthropus boisei: OH 5 anterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20boisei\/OH%205\">Paranthropus boisei: OH 5 inferior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20boisei\/OH%205\">Paranthropus boisei: OH 5 posterior view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong><em>Paranthropus robustus<\/em><\/strong> was the first taxon to be discovered within the genus in Kromdraai B by a schoolboy named Gert Terblanche; subsequent fossil discoveries were made by researcher Robert Broom in 1938 (Figure 9.22; Broom 1938a, 1938b, 1950), with the holotype specimen TM 1517 (Broom 1938a, 1938b, 1950; Hlazo 2018). <em>Paranthropus robustus<\/em> dates approximately from 2.0 mya to 1 mya and is the only taxon from the genus to be discovered in South Africa. Several of these fossils are fragmentary in nature, distorted, and not well preserved because they have been recovered from quarry breccia using explosives. <em>P. robustus<\/em> features are neither as \u201chyper-robust\u201d as <em>P. boisei<\/em> nor as ancestral as <em>P. aethiopicus<\/em>; instead, they have been described as being less derived, more general features that are shared with both East African species (e.g., the sagittal crest and zygomatic flaring; Rak 1983; Walker and Leakey 1988). Enamel hypoplasia is also common in this species, possibly because of instability in the development of large, thick-enameled dentition.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_305\" aria-describedby=\"caption-attachment-305\" style=\"width: 572px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-303 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.22.jpg\" alt=\"Four views of a beige-colored skull are shown on a black background.\" width=\"572\" height=\"619\" \/><figcaption id=\"caption-attachment-305\" class=\"wp-caption-text\">Figure 9.22: SK 48, a Paranthropus robustus specimen, had less derived, more general features that were not as robust as P. boisei and not as ancestral as P. aethiopicus. Credit: a. <a class=\"rId208\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId209\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 anterior view<\/a> by \u00a9<a class=\"rId210\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId211\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId212\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId213\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId214\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 superior view<\/a> by \u00a9<a class=\"rId215\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId216\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId217\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; c. <a class=\"rId218\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId219\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 inferior view<\/a> by \u00a9<a class=\"rId220\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId221\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId222\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; d. <a class=\"rId223\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\"><em>Paranthropus robustus<\/em><\/a><a class=\"rId224\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Paranthropus%20robustus\/SK%2048\">: SK 48 lateral left view<\/a> by \u00a9<a class=\"rId225\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId226\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId227\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Comparisons between Gracile and Robust Australopiths<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Comparisons between gracile and robust australopithecines may indicate different phylogenetic groupings or parallel evolution in several species. In general, the robust australopithecines have large temporalis (chewing) muscles, as indicated by flaring zygomatic arches, sagittal crests, and robust mandibles (jawbones). Their hind dentition is large (megadont), with low cusps and thick enamel. Within the gracile australopithecines, researchers have debated the relatedness of the species, or even whether these species should be lumped together to represent more variable or polytypic species. Often researchers will attempt to draw chronospecific trajectories, with one taxon said to evolve into another over time.<\/span><\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Special Topic: The Taung Child<\/span><\/h2>\n<figure id=\"attachment_304\" aria-describedby=\"caption-attachment-304\" style=\"width: 570px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-304 \" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/9.23.jpg\" alt=\"An ancient skull in anterior and lateral views. One view shows an imprint of the brain.\" width=\"570\" height=\"285\" \/><figcaption id=\"caption-attachment-304\" class=\"wp-caption-text\">Figure 9.23: The Taung Child has a nearly complete face, mandible, and partial endocranial cast. Credit: a. <em>A<\/em><a class=\"rId230\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\"><em>ustralopithecus africanus<\/em><\/a><a class=\"rId231\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\">: Taung 1 anterior view<\/a> by \u00a9<a class=\"rId232\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId233\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId234\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>; b. <a class=\"rId235\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\"><em>australopithecus africanus<\/em><\/a><a class=\"rId236\" href=\"https:\/\/efossils.org\/page\/boneviewer\/Australopithecus%20africanus\/Taung%201\">: Taung 1 lateral right view<\/a> by \u00a9<a class=\"rId237\" href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a class=\"rId238\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a class=\"rId239\" href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">The well-known fossil of a juvenile <em>Australopithecine<\/em>, the \u201cTaung Child,\u201d was the first early hominin evidence ever discovered and was the first to demonstrate our common human heritage in Africa (Figure 9.23; Dart 1925). The tiny facial skeleton and natural endocast were discovered in 1924 by a local quarryman in the North West Province in South Africa and were painstakingly removed from the surrounding cement-like breccia by Raymond Dart using his wife\u2019s knitting needles. When first shared with the scientific community in 1925, it was discounted as being nothing more than a young monkey of some kind. Prevailing biases of the time made it too difficult to contemplate that this small-brained hominin could have anything to do with our own history. The fact that it was discovered in Africa simply served to strengthen this bias.<\/span><\/p>\n<\/div>\n<p>&nbsp;<\/p>\n<h2><span style=\"color: #000000\">Early Tool Use and Technology<br \/>\n<\/span><\/h2>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Early Stone Age Technology (ESA)<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1378\">Early Stone Age (ESA)<\/a><\/strong> marks the beginning of recognizable technology made by our human ancestors. Stone-tool (or <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1380\"><strong>lithic<\/strong><\/a>) technology is defined by the fracturing of rocks and the manufacture of tools through a process called <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1382\"> <strong>knapping<\/strong><\/a>. The Stone Age lasted for more than 3 million years and is broken up into chronological periods called the Early (ESA), Middle (MSA), and Later Stone Ages (LSA). Each period is further broken up into a different <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1384\">techno-complex<\/a><\/strong>, a term encompassing multiple <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1386\">assemblages<\/a><\/strong> (collections of artifacts) that share similar traits in terms of artifact production and morphology. The ESA spanned the largest technological time period of human innovation from over 3 million years ago to around 300,000 years ago and is associated almost entirely with hominin species prior to modern <em>Homo sapiens. <\/em>As the ESA advanced, stone tool makers (known as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1390\"><strong>knappers<\/strong><\/a>) began to change the ways they detached <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1388\">flakes<\/a><\/strong> and eventually were able to shape artifacts into functional tools. These advances in technology go together with the developments in human evolution and cognition, dispersal of populations across the African continent and the world, and climatic changes.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">In order to understand the ESA, it is important to consider that not all assemblages are exactly the same within each techno-complex: one can have multiple phases and traditions at different sites (Lombard et al. 2012). However, there is an overarching commonality between them. Within stone tool assemblages, both flakes or <strong>cores<\/strong> (the rocks from which flakes are removed) are used as tools.<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1392\"> <strong>Large Cutting Tools (LCTs)<\/strong><\/a> are tools that are shaped to have functional edges. It is important to note that the information presented here is a small fraction of what is known about the ESA, and there are ongoing debates and discoveries within archaeology.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Currently, the oldest-known stone tools, which form the techno-complex the Lomekwian, date to 3.3 mya (Harmand et al. 2015; Toth 1985). They were found at a site called Lomekwi 3 in Kenya. This techno-complex is the most recently defined and pushed back the oldest-known date for lithic technology. There is only one known site thus far and, due to the age of the site, it is associated with species prior to <em>Homo<\/em>, such as <em>Kenyanthropus platyops.<\/em> Flakes were produced through indirect percussion, whereby the knappers held a rock and hit it against another rock resting on the ground. The pieces are very chunky and do not display the same fracture patterns seen in later techno-complexes. Lomekwian knappers likely aimed to get a sharp-edged piece on a flake, which would have been functional, although the specific function is currently unknown.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Stone tool use, however, is not only understood through the direct discovery of the tools. Cut marks on fossilized animal bones may illuminate the functionality of stone tools. In one controversial study in 2010, researchers argued that cut marks on a pair of animal bones from Dikika (Ethiopia), dated to 3.4 mya, were from stone tools. The discoverers suggested that they be more securely associated, temporally, with <em>Au. afarensis<\/em>. However, others have noted that these marks are consistent with teeth marks from crocodiles and other carnivores.<\/span><\/p>\n<figure style=\"width: 324px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image29-1.png\" alt=\"A technical line drawing of an Oldowan chopper.\" width=\"324\" height=\"275\" \/><figcaption class=\"wp-caption-text\">Figure 9.24: Some scholars believe that some genera explored in this chapter were capable of producing more complex stone tools (Oldowan). Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Olduwan_Industry_Chopper_2.jpeg\">Olduwan Industry Chopper 2<\/a> by Emmyanne29 is under a <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/legalcode\">CC0 1.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The Oldowan techno-complex is far more established in the scientific literature (Leakey 1971). It is called the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1394\">Oldowan<\/a><\/strong> because it was originally discovered in Olduvai Gorge, Tanzania, but the oldest assemblage is from Gona in Ethiopia, dated to 2.6 mya (Semaw 2000). The techno-complex is defined as a core and flake industry. Like the Lomekwian, there was an aim to get sharp-edged flakes, but this was achieved through a different production method. Knappers were able to actively hold or manipulate the core being knapped, which they could directly hit using a hammerstone. This technique is known as free-hand percussion, and it demonstrates an understanding of fracture mechanics. It has long been argued that the Oldowan hominins were skillful in tool manufacture.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Because Oldowan knapping requires skill, earlier researchers have attributed these tools to members of our genus, <em>Homo<\/em>. However, some have argued that these tools are in more direct association with hominins in the genera described in this chapter (Figure 9.24).<\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Invisible Tool Manufacture and Use<\/span><\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The vast majority of our understanding of these early hominins comes from fossils and reconstructed paleoenvironments. It is only from 3 mya when we can start \u201clooking into their minds\u201d and lifestyles by analyzing their manufacture and use of stone tools. However, the vast majority of tool use in primates (and, one can argue, in humans) is not with durable materials like stone. All of our extant great ape relatives have been observed using sticks, leaves, and other materials for some secondary purpose (to wade across rivers, to \u201cfish\u201d for termites, or to absorb water for drinking). It is possible that the majority of early hominin tool use and manufacture may be invisible to us because of this preservation bias.<\/span><\/p>\n<h2 class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Chapter Summary<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000\">The fossil record of our earliest hominin relatives has allowed paleoanthropologists to unpack some of the mysteries of our evolution. We now know that traits associated with bipedalism evolved before other \u201chuman-like\u201d traits, even though the first hominins were still very capable of arboreal locomotion. We also know that, for much of this time, hominin taxa were diverse in the way they looked and what they ate, and they were widely distributed across the African continent. And we know that the environments in which these hominins lived underwent many changes over this time during several warming and cooling phases.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Yet this knowledge has opened up many new mysteries. We still need to better differentiate some taxa. In addition, there are ongoing debates about why certain traits evolved and what they meant for the extinction of some of our relatives (like the robust australopiths). The capabilities of these early hominins with respect to tool use and manufacture is also still uncertain.<\/span><\/p>\n<h2 class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Hominin Species Summaries<br \/>\n<\/span><\/h2>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Sahelanthropus tchadensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">7 mya to 6 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Chad<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The initial discovery, made in 2001.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">360 cc average<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Smaller than in extant great apes; larger and pointier than in humans. Canines worn at the tips.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A short cranial base and a foramen magnum (hole in which the spinal cord enters the cranium) that is more humanlike in positioning; has been argued to indicate upright walking.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Currently little published postcranial material.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 0\">\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The extent to which this hominin was bipedal is currently heavily debated. If so, it would indicate an arboreal bipedal ancestor of hominins, not a knuckle-walker like chimpanzees.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Orrorin tugenensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">6 mya to 5.7 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Tugen Hills (Kenya)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Original discovery in 2000.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Smaller cheek teeth (molars and premolars) than even more recent hominins (i.e., derived), thick enamel, and reduced, but apelike, canines.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Not many found<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Fragmentary leg, arm, and finger bones have been found. Indicates bipedal locomotion.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Potential toolmaking capability based on hand morphology, but nothing found directly.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table5-R\" style=\"height: 0\">\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">This is the earliest species that clearly indicates adaptations for bipedal locomotion.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Ardipithecus kadabba<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">5.2 mya to 5.8 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Middle Awash (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Discovered by Yohannes Haile-Selassie in 1997.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Larger hind dentition than in modern chimpanzees. Thick enamel and larger canines than in later hominins.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A large hallux (big toe) bone indicates a bipedal \u201cpush off.\u201d<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table6-R\" style=\"height: 0\">\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table6-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Faunal evidence indicates a mixed grassland\/woodland environment.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Ardipithecus ramidus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">4.4 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Middle Awash region and Gona (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">A partial female skeleton nicknamed \u201cArdi\u201d (ARA-VP-6\/500) (found in 1994).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">300 cc to 350 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Little differences between the canines of males and females (small sexual dimorphism).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Midfacial projection, slightly prognathic. Cheekbones less flared and robust than in later hominins.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Ardi demonstrates a mosaic of ancestral and derived characteristics in the postcrania. For instance, an opposable big toe similar to chimpanzees (i.e., more ancestral), which could have aided in climbing trees effectively. However, the pelvis and hip show that she could walk upright (i.e., it is derived), supporting her hominin status.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">None directly associated<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table7-R\" style=\"height: 0\">\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table7-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">Over 110 specimens from Aramis<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus anamensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">4.2 mya to 3.8 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Turkana region (Kenya); Middle Awash (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A 2019 find from Ethiopia, named MRD.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">370 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Relatively large canines compared with more recent Australopithecines.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Projecting cheekbones and ancestral earholes.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lower limb bones (tibia and femur) indicate bipedality; arboreal features in upper limb bones (humerus) found.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table8-R\" style=\"height: 0\">\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table8-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Almost 100 specimens, representing over 20 individuals, have been found to date.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus afarensis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">3.9 mya to 2.9 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Afar Region, Omo, Maka, Fejej, and Belohdelie (Ethiopia); Laetoli (Tanzania); Koobi Fora (Kenya)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lucy (discovery: 1974), Selam (Dikika Child, discovery: 2000), Laetoli Footprints (discovery: 1976).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">380 cc to 430 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Reduced canines and molars relative to great apes but larger than in modern humans.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Prognathic face, facial features indicate relatively strong chewing musculature (compared with <em>Homo<\/em>) but less extreme than in <em>Paranthropus<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clear evidence for bipedalism from lower limb postcranial bones. Laetoli Footprints indicate humanlike walking. Dikika Child bones indicate retained ancestral arboreal traits in the postcrania.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">None directly, but close in age and proximity to controversial cut marks at Dikika and early tools in Lomekwi.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table9-R\" style=\"height: 0\">\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table9-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Au. afarensis<\/em> is one of the oldest and most well-known australopithecine species and consists of a large number of fossil remains.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus bahrelghazali<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">3.6 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Chad<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">\u201cAbel,\u201d the holotype (discovery: 1995).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table10-R\" style=\"height: 0\">\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table10-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Arguably within range of variation of <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus prometheus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">3.7 mya (debated)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">Sterkfontein (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">\u201cLittle Foot\u201d (StW 573) (discovery: 1994)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">408 cc (Little Foot estimate)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">Heavy anterior dental wear patterns, relatively large anterior dentition and smaller hind dentition, similar to <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;text-indent: 0pt\"><span style=\"color: #000000\">Relatively larger brain size, robust zygomatic arch, and a flatter midface.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">The initial discovery of four ankle bones indicated bipedality.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table11-R\" style=\"height: 0\">\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table11-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\">Highly debated new species designation.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus <\/em><em>deyiremada<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">3.5 mya to 3.3 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Woranso-Mille (Afar region, Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">First fossil mandible bones were discovered in 2011 in the Afar region of Ethiopia by Yohannes Haile-Selassie.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Smaller teeth with thicker enamel than seen in <em>Au. afarensis<\/em>, with a potentially hardier diet.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Larger mandible and more projecting cheekbones than in <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table12-R\" style=\"height: 0\">\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table12-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Contested species designation; arguably a member of <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Kenyanthopus<\/em><em> platyops<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">3.5 mya to 3.2 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Lake Turkana (Kenya)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">KNM\u2013WT 40000 (discovered 1999)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Difficult to determine but appears within the range of <em>Australopithecus afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Small molars\/dentition (<em>Homo<\/em>-like characteristic)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Flatter (i.e., orthognathic) face<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Some have associated the earliest tool finds from Lomekwi, Kenya, temporally (3.3 mya) and in close geographic proximity to this species\/specimen.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table13-R\" style=\"height: 0\">\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table13-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Taxonomic placing of this species is quite divided. The discoverers have argued that this species is ancestral to <em>Homo<\/em>, in particular to <em>Homo <\/em><em>ruldolfensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Australopithecus africanus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">3.3 mya to 2.1 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Sterkfontein, Taung, Makapansgat, Gladysvale (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Taung Child (discovery in 1994), \u201cMrs. Ples\u201d (discover in 1947), Little Foot (arguable; discovery in 1994).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">400 cc to 500 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Smaller teeth (derived) relative to <em>Au. afarensis<\/em>. Small canines with no diastema.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">A rounder skull compared with <em>Au. afarensis<\/em> in East Africa. A sloping face (ancestral).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Similar postcranial evidence for bipedal locomotion (derived pelvis) with retained arboreal locomotion, e.g., curved phalanges (fingers), as seen in <em>Au. afarensis.<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">None with direct evidence.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table14-R\" style=\"height: 0\">\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table14-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">A 2015 study noted that the trabecular bone morphology of the hand was consistent with forceful tool manufacture and use, suggesting potential early tool abilities.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Australopithecus garhi<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.5 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Middle Awash (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">450 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Larger hind dentition than seen in other gracile Australopithecines.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A femur of a fragmentary partial skeleton, argued to belong to <em>Au. garhi<\/em>, indicates this species may be longer-limbed than <em>Au. afarensis<\/em>, although still able to move arboreally.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Crude stone tools resembling Oldowan (described later) have been found in association with <em>Au. garhi<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table15-R\" style=\"height: 0\">\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table15-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">This species is not well documented or understood and is based on only a few fossil specimens.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Paranthropus aethiopicus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.7 mya to 2.3 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">West Turkana (Kenya); Laetoli (Tanzania); Omo River Basin (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The \u201cBlack Skull\u201d (KNM\u2013WT 17000) (discovery 1985).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain Size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">410 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>P. aethiopicus<\/em> has the shared derived traits of large flat premolars and molars, although few teeth have been found.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Large flaring zygomatic arches for accommodating large chewing muscles (the temporalis muscle), a sagittal crest for increased muscle attachment of the chewing muscles to the skull, and a robust mandible and supraorbital torus (brow ridge).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A proximal tibia indicates bipedality and similar size to <em>Au. afarensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table16-R\" style=\"height: 0\">\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table16-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 1.5pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">The \u201cBlack Skull\u201d is so called because of the mineral manganese that stained it black during fossilization.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Paranthropus boisei<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.4 mya to 1.4 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Koobi Fora, West Turkana, and Chesowanja (Kenya); Malema-Chiwondo (Malawi), Olduvai Gorge and Peninj (Tanzania); and Omo River basin and Konso (Ethiopia)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">\u201cZinj,\u201d or sometimes \u201cNutcracker Man\u201d (OH5), in 1959 by Mary Leakey. The Peninj mandible from Tanzania, found in 1964 by Kimoya Kimeu.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">500 cc to 550 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Very large, flat posterior dentition (largest of all hominins currently known). Much smaller anterior dentition. Very thick dental enamel.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Indications of very large chewing muscles (e.g., flaring zygomatic arches and a large sagittal crest).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Evidence for high variability and sexual dimorphism, with estimates of males at 1.37 meters tall and females at 1.24 meters.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Richard Leakey and Bernard Wood have both suggested that<em> P. boisei<\/em> could have made and used stone tools. Tools dated to 2.5 mya in Ethiopia have been argued to possibly belong to this species.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table17-R\" style=\"height: 0\">\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table17-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Despite the cranial features of <em>P. boisei<\/em> indicating a tough diet of tubers, nuts, and seeds, isotopes indicate a diet high in C4 foods (e.g., grasses, such as sedges). This differs from what is seen in<em> P. robustus<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\"><em>Australopithecus sediba<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">1.97 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Malapa Fossil Site (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Karabo (MH1) (discovery in 2008)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">420 cc to 450 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Small dentition with Australopithecine cusp-spacing.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Small brain size (<em>Australopithecus<\/em>-like) but gracile mandible (<em>Homo<\/em>-like).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Scientists have interpreted this mixture of traits (such as a robust ankle but evidence for an arch in the foot) as a transitional phase between a body previously adapted to arborealism (tree climbing, particularly in evidence from the bones of the wrist) to one that adapted to bipedal ground walking.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">None of direct association, but some have argued that a modern hand morphology (shorter fingers and a longer thumb) means that adaptations to tool manufacture and use may be present in this species.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table18-R\" style=\"height: 0\">\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table18-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">It was first discovered through a clavicle bone in 2008 by nine-year-old Matthew Berger, son of paleoanthropologist Lee Berger.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left\">\n<table style=\"width: 450pt\">\n<tbody>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>Paranthropus robustus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">2.3 mya to 1 mya<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kromdraai B, Swartkrans, Gondolin, Drimolen, and Coopers Cave (South Africa)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">SK48 (original skull)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Brain <\/strong><strong>s<\/strong><strong>ize<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">410 cc to 530 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Large posterior teeth with thick enamel, consistent with other Robust Australopithecines. Enamel hypoplasia is also common in this species, possibly because of instability in the development of large, thick enameled dentition.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><em>P. robustus<\/em> features are neither as \u201chyper-robust\u201d as <em>P. boisei<\/em> or as ancestral in features as <em>P. aethiopicus<\/em>. They have been described as less derived, more general features that are shared with both East African species (e.g., the sagittal crest and zygomatic flaring).<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Reconstructions indicate sexual dimorphism.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table19-R\" style=\"height: 0\">\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table19-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Several of these fossils are fragmentary in nature, distorted, and not well preserved, because they have been recovered from quarry breccia using explosives.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\"><strong><span style=\"color: #000000\">Review Questions<br \/>\n<\/span><\/strong><\/h2>\n<ul>\n<li class=\"import-Normal\"><span style=\"color: #000000\">What is the difference between a \u201cderived\u201d versus an \u201cancestral\u201d trait? Give an example of both, seen in <em>Au. afarensis<\/em>.<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">Which of the paleoenvironment hypotheses have been used to describe early hominin diversity, and which have been used to describe bipedalism?<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">Which anatomical features for bipedalism do we see in early hominins?<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">Describe the dentition of gracile and robust australopithecines. What might these tell us about their diets?<\/span><\/li>\n<li class=\"import-Normal\"><span style=\"color: #000000\">List the hominin species argued to be associated with stone tool technologies. Are you convinced of these associations? Why\/why not?<\/span><\/li>\n<\/ul>\n<\/div>\n<h2><span style=\"color: #000000\">Key Terms<\/span><\/h2>\n<p><span style=\"color: #000000\"><strong>Arboreal:<\/strong> Related to trees or woodland.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Aridification:<\/strong> Becoming increasingly arid or dry, as related to the climate or environment.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Aridity Hypothesis:<\/strong> The hypothesis that long-term aridification and expansion of savannah biomes were drivers in diversification in early hominin evolution.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Assemblage:<\/strong> A collection demonstrating a pattern. Often pertaining to a site or region.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Bipedalism:<\/strong> The locomotor ability to walk on two legs.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Breccia:<\/strong> Hard, calcareous sedimentary rock.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Canines:<\/strong> The pointy teeth just next to the incisors, in the front of the mouth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cheek teeth:<\/strong> Or hind dentition (molars and premolars).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Chronospecies:<\/strong> Species that are said to evolve into another species, in a linear fashion, over time.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Clade:<\/strong> A group of species or taxa with a shared common ancestor.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cladistics:<\/strong> The field of grouping organisms into those with shared ancestry.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Context:<\/strong> As pertaining to palaeoanthropology, this term refers to the place where an artifact or fossil is found.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cores:<\/strong> The remains of a rock that has been flaked or knapped.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Cusps:<\/strong> The ridges or \u201cbumps\u201d on the teeth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Dental formula:<\/strong> A technique to describe the number of incisors, canines, premolars, and molars in each quadrant of the mouth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Derived traits:<\/strong> Newly evolved traits that differ from those seen in the ancestor.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Diastema:<\/strong> A tooth gap between the incisors and canines.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Early Stone Age (ESA):<\/strong> The earliest-described archaeological period in which we start seeing stone-tool technology.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>East African Rift System (EARS):<\/strong> This term is often used to refer to the Rift Valley, expanding from Malawi to Ethiopia. This active geological structure is responsible for much of the visibility of the paleoanthropological record in East Africa.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Enamel:<\/strong> The highly mineralized outer layer of the tooth.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Encephalization:<\/strong> Expansion of the brain.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Extant:<\/strong> Currently living\u2014i.e., not extinct.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Fallback foods:<\/strong> Foods that may not be preferred by an animal (e.g., foods that are not nutritionally dense) but that are essential for survival in times of stress or scarcity.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Fauna:<\/strong> The animals of a particular region, habitat, or geological period.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Faunal assemblages:<\/strong> Collections of fossils of the animals found at a site.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Faunal turnover:<\/strong> The rate at which species go extinct and are replaced with new species.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Flake:<\/strong> The piece knocked off of a stone core during the manufacture of a tool, which may be used as a stone tool.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Flora:<\/strong> The plants of a particular region, habitat, or geological period.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Folivorous:<\/strong> Foliage-eating.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Foramen magnum:<\/strong> The large hole (foramen) at the base of the cranium, through which the spinal cord enters the skull.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Fossil:<\/strong> The remains or impression of an organism from the past.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Frugivorous:<\/strong> Fruit-eating.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Generalist:<\/strong> A species that can thrive in a wide variety of habitats and can have a varied diet.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Glacial:<\/strong> Colder, drier periods during an ice age when there is more ice trapped at the poles.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Gracile:<\/strong> Slender, less rugged, or pronounced features.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hallux:<\/strong> The big toe.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Holotype:<\/strong> A single specimen from which a species or taxon is described or named.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hominin:<\/strong> A primate category that includes humans and our fossil relatives since our divergence from extant great apes.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Honing P3:<\/strong> The mandibular premolar alongside the canine (in primates, the P3), which is angled to give space for (and sharpen) the upper canines.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hyper-robust:<\/strong> Even more robust than considered normal in the Paranthropus genus.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Hypodigm:<\/strong> A sample (here, fossil) from which researchers extrapolate features of a population.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Incisiform:<\/strong> An adjective referring to a canine that appears more incisor-like in morphology.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Incisors:<\/strong> The teeth in the front of the mouth, used to bite off food.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Interglacial:<\/strong> A period of milder climate in between two glacial periods.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Isotopes:<\/strong> Two or more forms of the same element that contain equal numbers of protons but different numbers of neutrons, giving them the same chemical properties but different atomic masses.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Knappers:<\/strong> The people who fractured rocks in order to manufacture tools.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Knapping:<\/strong> The fracturing of rocks for the manufacture of tools.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Large Cutting Tool (LCT):<\/strong> A tool that is shaped to have functional edges.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Last Common Ancestor (LCA):<\/strong> The hypothetical final ancestor (or ancestral population) of two or more taxa before their divergence.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Lithic:<\/strong> Relating to stone (here to stone tools).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Lumbar lordosis:<\/strong> The inward curving of the lower (lumbar) parts of the spine. The lower curve in the human S-shaped spine.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Lumpers:<\/strong> Researchers who prefer to lump variable specimens into a single species or taxon and who feel high levels of variation is biologically real.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Megadont:<\/strong> An organism with extremely large dentition compared with body size.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Metacarpals:<\/strong> The long bones of the hand that connect to the phalanges (finger bones).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Molars:<\/strong> The largest, most posterior of the hind dentition.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Monophyletic:<\/strong> A taxon or group of taxa descended from a common ancestor that is not shared with another taxon or group.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Morphology:<\/strong> The study of the form or size and shape of things; in this case, skeletal parts.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Mosaic evolution:<\/strong> The concept that evolutionary change does not occur homogeneously throughout the body in organisms.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Obligate bipedalism:<\/strong> Where the primary form of locomotion for an organism is bipedal.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Occlude:<\/strong> When the teeth from the maxilla come into contact with the teeth in the mandible.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Oldowan:<\/strong> Lower Paleolithic, the earliest stone tool culture.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Orthognathic:<\/strong> The face below the eyes is relatively flat and does not jut out anteriorly.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Paleoanthropologists:<\/strong> Researchers that study human evolution.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Paleoenvironment:<\/strong> An environment from a period in the Earth\u2019s geological past.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Parabolic:<\/strong> Like a parabola (parabola-shaped).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Phalanges:<\/strong> Long bones in the hand and fingers.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Phylogenetics:<\/strong> The study of phylogeny.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Phylogeny:<\/strong> The study of the evolutionary relationships between groups of organisms.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Pliocene:<\/strong> A geological epoch between the Miocene and Pleistocene.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Polytypic:<\/strong> In reference to taxonomy, having two or more group variants capable of interacting and breeding biologically but having morphological population differences.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Postcranium:<\/strong> The skeleton below the cranium (head).<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Premolars:<\/strong> The smallest of the hind teeth, behind the canines.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Procumbent:<\/strong> In reference to incisors, tilting forward.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Prognathic:<\/strong> In reference to the face, the area below the eyes juts anteriorly.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Quaternary Ice Age:<\/strong> The most recent geological time period, which includes the Pleistocene and Holocene Epochs and which is defined by the cyclicity of increasing and decreasing ice sheets at the poles.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Relative dating:<\/strong> Dating techniques that refer to a temporal sequence (i.e., older or younger than others in the reference) and do not estimate actual or absolute dates.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Robust:<\/strong> Rugged or exaggerated features.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Site:<\/strong> A place in which evidence of past societies\/species\/activities may be observed through archaeological or paleontological practice.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Specialist:<\/strong> A specialist species can thrive only in a narrow range of environmental conditions or has a limited diet.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Splitters:<\/strong> Researchers who prefer to split a highly variable taxon into multiple groups or species.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Taxa:<\/strong> Plural of taxon, a taxonomic group such as species, genus, or family.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Taxonomy:<\/strong> The science of grouping and classifying organisms.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Techno-complex:<\/strong> A term encompassing multiple assemblages that share similar traits in terms of artifact production and morphology.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Thermoregulation:<\/strong> Maintaining body temperature through physiologically cooling or warming the body.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Ungulates:<\/strong> Hoofed mammals\u2014e.g., cows and kudu.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Volcanic tufts:<\/strong> Rock made from ash from volcanic eruptions in the past.<\/span><\/p>\n<p><span style=\"color: #000000\"><strong>Valgus knee:<\/strong> The angle of the knee between the femur and tibia, which allows for weight distribution to be angled closer to the point above the center of gravity (i.e., between the feet) in bipeds.<\/span><\/p>\n<h2 class=\"import-Normal\"><strong><span style=\"color: #000000\">About the Authors<br \/>\n<\/span><\/strong><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image31.jpg\" alt=\"A woman with short blonde hair smiles at the camera.\" width=\"311\" height=\"311\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Kerryn Warren, Ph.D.<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Grad Coach International, <a class=\"rId245\" style=\"color: #000000\" href=\"mailto:kerryn.warren@gmail.com\">kerryn.warren@gmail.com<\/a><\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kerryn Warren is a dissertation coach at Grad Coach International and is passionate about stimulating research thinking in students of all levels. She has lectured on multiple topics, including archaeology and human evolution, with her research and science communication interests including hybridization in the hominin fossil record (stemming from research from her Ph.D.) and understanding how evolution is taught in South African schools. She also worked as one of the \u201cUnderground Astronauts,\u201d selected to excavate <em>Homo naledi <\/em>remains from the Rising Star Cave System in the Cradle of Humankind.<\/span><\/p>\n<p>&nbsp;<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image11-2.jpg\" alt=\"A woman with short brown hair smiles at the camera.\" width=\"312\" height=\"306\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">K. Lindsay Hunter, M.A., Ph.D. candidate<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">CARTA, k.lindsay.hunter@gmail.com<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lindsay Hunter is a trained palaeoanthropologist who uses her more than 15 years of experience to make sense of the distant past of our species to build a better future. She received her master\u2019s degree in biological anthropology from the University of Iowa and is completing her Ph.D. in archaeology at the University of the Witwatersrand in Johannesburg, South Africa. She has studied fossil and human bone collections across five continents with major grant support from the National Science Foundation (United States) and the Wenner-Gren Foundation for Anthropological Research. As a National Geographic Explorer, Lindsay developed and managed the National Geographic\u2013sponsored Umsuka Public Palaeoanthropology Project in the Cradle of Humankind World Heritage Site (CoH WHS) in South Africa from within Westbury Township, Johannesburg, between 2016\u20132019. She currently serves as the Community Engagement &amp; Advancement Director for CARTA: The UC San Diego\/Salk Institute Center for Academic Research and Training in Anthropogeny in La Jolla, California.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image25-2.jpg\" alt=\"A woman with black hair stands in a hole in the ground.\" width=\"295\" height=\"339\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Navashni Naidoo, M.Sc.<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">University of Cape Town, nnaidoo2@illinois.edu<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Navashni Naidoo is a researcher at Nelson Mandela University, lecturing on physical geology. She completed her Master\u2019s in Science in Archaeology in 2017 at the University of Cape Town. Her research interests include developing paleoenvironmental proxies suited to the African continent, behavioral ecology, and engaging with community-driven archaeological projects. She has excavated at Stone Age sites across Southern Africa and East Africa. Navashni is currently pursuing a PhD in the Department of Anthropology at the University of Illinois.<\/span><\/p>\n<p>&nbsp;<\/p>\n<p>&nbsp;<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image47.jpg\" alt=\"A man with black hair and dark brown eyes looks at the camera. \" width=\"294\" height=\"294\" \/><\/span><\/p>\n<h3 class=\"import-Normal\"><strong><span style=\"color: #000000\">Silindokuhle Mavuso, M.Sc.<\/span><\/strong><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000\">University of Witwatersrand, S.muvaso@ru.ac.za<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Silindokuhle has always been curious about the world around him and how it has been shaped. He is a lecturer at Rhodes University of Witwatersrand (Wits), and conducts research on palaeoenvironmental reconstruction and change of the northeastern Turkana Basin\u2019s Pleistocene sequence. Silindokuhle began his education with a B.Sc. (Geology, Archaeology, and Environmental and Geographical Sciences) from the University of Cape Town before moving to Wits for a B.Sc. Honors (geology and paleontology) and M.Sc. in geology. He is currently concluding his PhD Studies. During this time, he has gained more training as a Koobi Fora Fieldschool fellow (Kenya) as well as an Erasmus Mundus scholar (France). Silindokuhle is a Plio-Pleistocene geologist with a specific focus on identifying and explaining past environments that are associated with early human life and development through time. He is interested in a wide range of disciplines such as micromorphology, sedimentology, geochemistry, geochronology, and sequence stratigraphy. He has worked with teams from significant eastern and southern African hominid sites including Elandsfontein, Rising Star, Sterkfontein, Gondolin, Laetoli, Olduvai, and Koobi Fora.<\/span><\/p>\n<h2 class=\"import-Normal\"><strong><span style=\"color: #000000\">For Further Exploration<br \/>\n<\/span><\/strong><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/humanorigins.si.edu\/evidence\">The Smithsonian Institution website<\/a> hosts descriptions of fossil species, an interactive timeline, and much more.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.maropeng.co.za\/content\/page\/human-evolution\">The Maropeng Museum website<\/a> hosts a wealth of information regarding South African Fossil Bearing sites in the Cradle of Humankind<strong>.<\/strong><\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/perot-museum.imgix.net\/2019-08-naledi-sediba-quick-comparison.pdf\">This quick comparison between <em>Homo naledi<\/em> and <em>Australopithecus sediba<\/em><\/a> from the Perot Museum.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.dropbox.com\/s\/l1d2hv42psj21y9\/Braided%20Stream-1920.mp4?dl=0\">This explanation of the braided stream<\/a> by the Perot Museum.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.hetmp.com\/\">A collation of 3-D files for visualizing<\/a> (or even 3-D printing) for homes, schools, and universities.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\"><a href=\"https:\/\/www.pbslearningmedia.org\/resource\/tdc02.sci.life.evo.lp_humanevo\/human-evolution.\">PBS learning materials<\/a>, including videos and diagrams of the Laetoli footprints, bipedalism, and fossils.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">A wealth of <a href=\"https:\/\/australianmuseum.net.au\/learn\/science\/human-evolution\/\">information from the Australian Museum website<\/a>, including species descriptions, family trees, and explanations of bipedalism and diet<strong>.<\/strong><\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\"><strong>References<\/strong><\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Alemseged, Zeresenay, Fred Spoor, William H. Kimbel, Ren\u00e9 Bobe, Denis Geraads, Denn\u00e9 Reed, and Jonathan G. Wynn. 2006. \u201cA Juvenile Early Hominin Skeleton from Dikika, Ethiopia.\u201d <em>Nature<\/em> 443 (7109): 296\u2013301.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Asfaw, Berhane, Tim White, Owen Lovejoy, Bruce Latimer, Scott Simpson, and Gen Suwa. 1999. \u201c<em>Australopithecus garhi<\/em>: A New Species of Early Hominid from Ethiopia.\u201d <em>Science<\/em> 284 (5414): 629\u2013635.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Behrensmeyer, Anna K., Nancy E. Todd, Richard Potts, and Geraldine E. McBrinn. 1997. \u201cLate Pliocene Faunal Turnover in the Turkana Basin, Kenya, and Ethiopia.\u201d <em>Science<\/em> 278 (5343): 637\u2013640.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Berger, Lee R., Darryl J. De Ruiter, Steven E. Churchill, Peter Schmid, Kristian J. Carlson, Paul HGM Dirks, and Job M. Kibii. 2010. \u201c<em>Australopithecus sediba<\/em>: A New Species of <em>Homo<\/em>-like Australopith from South Africa.\u201d <em>Science<\/em> 328 (5975): 195\u2013204.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Bobe, Ren\u00e9, and Anna K. Behrensmeyer. 2004. \u201cThe Expansion of Grassland Ecosystems in Africa in Relation to Mammalian Evolution and the Origin of the Genus <em>Homo<\/em>.\u201d <em>Palaeogeography, Palaeoclimatology, Palaeoecology<\/em> 207 (3\u20134): 399\u2013420.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Brain, C. K. 1967. \u201cThe Transvaal Museum's Fossil Project at Swartkrans.\u201d <em>South African Journal of Science<\/em> 63 (9): 378\u2013384.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1938a. \u201cMore Discoveries of Australopithecus.\u201d <em>Nature<\/em> 141 (1): 828\u2013829.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1938b. \u201cThe Pleistocene Anthropoid Apes of South Africa.\u201d <em>Nature<\/em> 142 (3591): 377\u2013379.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1947. \u201cDiscovery of a New Skull of the South African Ape-Man, Plesianthropus.\u201d <em>Nature<\/em> 159 (4046): 672.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Broom, R. 1950. \u201cThe Genera and Species of the South African Fossil Ape-Man.\u201d <em>American Journal of Physical Anthropology<\/em> 8 (1): 1\u201314.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Brunet, Michel, Alain Beauvilain, Yves Coppens, Emile Heintz, Aladji HE Moutaye, and David Pilbeam. 1995. \u201cThe First Australopithecine 2,500 Kilometers West of the Rift Valley (Chad).\u201d <em>Nature<\/em> 378 (6554): 275\u2013273.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Cerling, Thure E., Jonathan G. Wynn, Samuel A. Andanje, Michael I. Bird, David Kimutai Korir, Naomi E. Levin, William Mace, Anthony N. Macharia, Jay Quade, and Christopher H. Remien. 2011. \u201cWoody Cover and Hominin Environments in the Past 6 Million Years.\u201d <em>Nature<\/em> 476, no. 7358 (2011): 51-56..<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clarke, Ronald J. 1998. \u201cFirst Ever Discovery of a Well-Preserved Skull and Associated Skeleton of <em>Australopithecus<\/em>.\u201d <em>South African Journal of Science<\/em> 94 (10): 460\u2013463.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Clarke, Ronald J. 2013. \u201cAustralopithecus from Sterkfontein Caves, South Africa.\u201d In <em>The Paleobiology of Australopithecus<\/em>, edited by K. E. Reed, J. G. Fleagle, and R. E. Leakey, 105\u2013123. Netherlands: Springer.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clarke, Ronald J., and Kathleen Kuman. 2019. \u201cThe Skull of StW 573, a 3.67 Ma Australopithecus Prometheus Skeleton from Sterkfontein Caves, South Africa.\u201d <em>Journal of <\/em><em>H<\/em><em>uman <\/em><em>E<\/em><em>volution<\/em> 134: 102634.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Clarke, R. J., and P. V. Tobias. 1995. \u201cSterkfontein Member 2 Foot Bones of the Oldest South African Hominid.\u201d <em>Science<\/em> 269 (5223): 521\u2013524.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Constantino, P. J., and B. A. Wood. 2004. \u201cParanthropus Paleobiology\u201d. In <em>Miscelanea en <\/em><em>Homenae<\/em><em> a Emiliano Aguirre<\/em><em>,<\/em> <em>v<\/em><em>olumen III: Paleoantropologia<\/em>, edited by E. G. P\u00e9rez and S. R. Jara, 136\u2013151. Alcal\u00e1 de Henares: Museo Arqueologico Regional.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Constantino, P. J., and B. A. Wood. 2007. \u201cThe Evolution of Zinjanthropus boisei.\u201d <em>Evolutionary Anthropology: <\/em><em>Issues, News, and Reviews<\/em> 16 (2): 49\u201362.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Dart, Raymond A. 1925. \u201cAustralopithecus africanus, the Man-Ape of South Africa.\u201d <em>Nature<\/em> 115: 195\u2013199.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Darwin, Charles. 1871. <em>The Descent of Man: And Selection in Relation to Sex<\/em>. London: J. Murray.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Daver, Guillaume, F. Guy, Hassane Ta\u00efsso Mackaye, Andossa Likius, J-R. Boisserie, Abderamane Moussa, Laurent Pallas, Patrick Vignaud, and N\u00e9koulnang D. Clarisse. 2022. \"Postcranial Evidence of Late Miocene Hominin Bipedalism in Chad.\" <em>Nature<\/em> 609 (7925): 94\u2013100.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Heinzelin, Jean de, J. Desmond Clark, Tim White, William Hart, Paul Renne, Giday WoldeGabriel, Yonas Beyene, and Elisabeth Vrba. 1999. \u201cEnvironment and Behavior of 2.5-Million-Year-Old Bouri Hominids.\u201d <em>Science<\/em> 284 (5414): 625\u2013629.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">DeMenocal, Peter B. D. 2004. \u201cAfrican Climate Change and Faunal Evolution during the Pliocene\u2013Pleistocene.\u201d <em>Earth and Planetary Science Letters<\/em> 220 (1\u20132): 3\u201324.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">DeMenocal, Peter B. D. and J. Bloemendal, J. 1995. \u201cPlio-Pleistocene Climatic Variability in Subtropical Africa and the Paleoenvironment of Hominid Evolution: A Combined Data-Model Approach.\u201d In <em>Paleoclimate and Evolution, with Emphasis on Human Origins<\/em>, edited by E. S. Vrba, G. H. Denton, T. C. Partridge, and L. H. Burckle, 262\u2013288. New Haven: Yale University Press.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Dirks, Paul HGM, Job M. Kibii, Brian F. Kuhn, Christine Steininger, Steven E. Churchill, Jan D. Kramers, Robyn Pickering, Daniel L. Farber, Anne-Sophie M\u00e9riaux, Andy I. R. Herries, Geoffrey C. P. King, And Lee R. Berger. 2010. \u201cGeological Setting and Age of <em>Australopithecus sediba<\/em> from Southern Africa.\u201d <em>Science<\/em> 328 (5975): 205\u2013208.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Faith, J. Tyler, and Anna K. Behrensmeyer. 2013. \u201cClimate Change and Faunal Turnover: Testing the Mechanics of the Turnover-Pulse Hypothesis with South African Fossil Data.\u201d <em>Paleobiology<\/em> 39 (4): 609\u2013627.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Grine, Frederick E. 1988. \u201cNew Craniodental Fossils of <em>Paranthropus<\/em> from the Swartkrans Formation and Their Significance in \u2018Robust\u2019 Australopithecine Evolution.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by F. E. Grine, 223\u2013243. New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Grine, Frederick E., Carrie S. Mongle, John G. Fleagle, and Ashley S. Hammond. 2022. \"The Taxonomic Attribution of African Hominin Postcrania from the Miocene through the Pleistocene: Associations and Assumptions.\" <em>Journal of Human Evolution<\/em> 173: 103255.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Haile-Selassie, Yohannes, Luis Gibert, Stephanie M. Melillo, Timothy M. Ryan, Mulugeta Alene, Alan Deino, Naomi E. Levin, Gary Scott, and Beverly Z. Saylor. 2015. \u201cNew Species from Ethiopia Further Expands Middle Pliocene Hominin Diversity.\u201d <em>Nature<\/em> 521 (7553): 432\u2013433.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Haile-Selassie, Yohannes, Stephanie M. Melillo, Antonino Vazzana, Stefano Benazzi, and Timothy M. Ryan. 2019. \u201cA 3.8-Million-Year-Old Hominin Cranium from Woranso-Mille, Ethiopia.\u201d <em>Nature<\/em> 573 (7773): 214-219.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Harmand, Sonia, Jason E. Lewis, Craig S. Feibel, Christopher J. Lepre, Sandrine Prat, Arnaud Lenoble, Xavier Bo\u00ebs et al. 2015. \u201c3.3-Million-Year-Old Stone Tools from Lomekwi3, West Turkana, Kenya.\u201d <em>Nature<\/em> 521 (7552): 310\u2013316.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Hay, Richard L. 1990. \u201cOlduvai Gorge: A Case History in the Interpretation of Hominid Paleoenvironments.\u201d In <em>East Africa: Establishment of a Geologic Framework for Paleoanthropology<\/em>, edited by L. Laporte, 23\u201337<em>.<\/em> Boulder: Geological Society of America.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Hay, Richard L., and Mary D. Leakey. 1982. \u201cThe Fossil Footprints of Laetoli.\u201d <em>Scientific American<\/em> 246 (2): 50\u201357.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Hlazo, Nomawethu. 2015. \u201cParanthropus: Variation in Cranial Morphology.\u201d Honours thesis, Archaeology Department, University of Cape Town, Cape Town.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Hlazo, Nomawethu. 2018. \u201cVariation and the Evolutionary Drivers of Diversity in the Genus <em>Paranthropus<\/em>.\u201d Master\u2019s thesis, Archaeology Department, University of Cape Town, Cape Town.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Johanson, D. C., T. D. White, and Y. Coppens. 1978. \u201cA New Species of the Genus <em>Australopithecus<\/em> (Primates: Hominidae) from the Pliocene of East Africa.\u201d <em>Kirtlandia<\/em> 28: 1\u201314.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kimbel, William H. 2015. \u201cThe Species and Diversity of Australopiths.\u201d In <em>Handbook of Paleoanthropology<\/em>, 2nd ed., edited by T. Hardt, 2071\u20132105. Berlin: Springer.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Kimbel, William H., and Lucas K. Delezene. 2009. \u201c\u2018Lucy\u2019 Redux: A Review of Research on <em>Australopithecus afarensis<\/em>.\u201d <em>American <\/em><em>J<\/em><em>ournal of <\/em><em>P<\/em><em>hysical <\/em><em>A<\/em><em>nthropology<\/em> 140 (S49): 2\u201348.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Kingston, John D. 2007. \u201cShifting Adaptive Landscapes: Progress and Challenges in Reconstructing Early Hominid Environments.\u201d <em>American Journal of Physical Anthropology<\/em> 134 (S45): 20\u201358.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Kingston, John D., and Terry Harrison. 2007. \u201cIsotopic Dietary Reconstructions of Pliocene Herbivores at Laetoli: Implications for Early Hominin Paleoecology.\u201d <em>Palaeogeography, Palaeoclimatology, Palaeoecology<\/em> 243 (3\u20134): 272\u2013306.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Leakey, Louis S. B. 1959. \u201cA New Fossil Skull from Olduvai.\u201d <em>Nature<\/em> 184 (4685): 491\u2013493.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Leakey, Mary 1971. <em>Olduvai Gorge<\/em>, Vol. 3. Cambridge: Cambridge University Press.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Leakey, Mary D., and Richard L. Hay. 1979. \u201cPliocene Footprints in the Laetoli Beds at Laetoli, Northern Tanzania.\u201d <em>Nature<\/em> 278 (5702): 317\u2013323.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Leakey, Meave G., Craig S. Feibel, Ian McDougall, and Alan Walker. 1995. \u201cNew Four\u2013Million-Year-Old Hominid Species from Kanapoi and Allia Bay, Kenya.\u201d <em>Nature<\/em> 376 (6541): 565\u2013571.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Meave G., Fred Spoor, Frank H. Brown, Patrick N. Gathogo, Christopher Kiarie, Louise N. Leakey, and Ian McDougall. 2001. \u201cNew Hominin Genus from Eastern Africa Shows Diverse Middle Pliocene Lineages.\u201d <em>Nature<\/em> 410 (6827): 433\u2013440.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lebatard, Anne-Elisabeth, Didier L. Bourl\u00e8s, Philippe Duringer, Marc Jolivet, R\u00e9gis Braucher, Julien Carcaillet, Mathieu Schuster et al. 2008. \u201cCosmogenic Nuclide Dating of <em>Sahelanthropus tchadensis<\/em> and <em>Australopithecus bahrelghazali<\/em>: Mio-Pliocene Hominids from Chad.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 105 (9): 3226\u20133231.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lee-Thorp, Julia. 2011. \u201cThe Demise of \u2018Nutcracker Man.\u2019\u201d <em>Proceedings of the National Academy of Sciences<\/em> 108 (23): 9319\u20139320.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Lombard, Marlize, L. Y. N. Wadley, Janette Deacon, Sarah Wurz, Isabelle Parsons, Moleboheng Mohapi, Joane Swart, and Peter Mitchell. 2012. \u201cSouth African and Lesotho Stone Age Sequence Updated.\u201d <em>The South African Archaeological Bulletin<\/em> 67 (195): 123\u2013144.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Maslin, Mark A., Chris M. Brierley, Alice M. Milner, Susanne Shultz, Martin H. Trauth, and Katy E. Wilson. 2014. \u201cEast African Climate Pulses and Early Human Evolution.\u201d <em>Quaternary Science Reviews<\/em> 101: 1\u201317.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">McHenry, Henry M. 2009. \u201cHuman Evolution.\u201d In <em>Evolution: The First Four Billion Years<\/em>, edited by M. Ruse and J. Travis, 256\u2013280. Cambridge: The Belknap Press of Harvard University Press..<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Patterson, Bryan, and William W. Howells. 1967. \u201cHominid Humeral Fragment from Early Pleistocene of Northwestern Kenya.\u201d <em>Science<\/em> 156 (3771): 64\u201366.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Pickering, Robyn, and Jan D. Kramers. 2010. \u201cRe-appraisal of the Stratigraphy and Determination of New U-Pb Dates for the Sterkfontein Hominin Site.\u201d <em>Journal of Human Evoluti<\/em><em>on<\/em> 59 (1): 70\u201386.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Potts, Richard. 1998. \u201cEnvironmental Hypotheses of Hominin Evolution.\u201d <em>American Journal of Physical Anthropology<\/em> 107 (S27): 93\u2013136.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Potts, Richard. 2013. \u201cHominin Evolution in Settings of Strong Environmental Variability.\u201d <em>Quaternary Science Reviews<\/em> 73: 1\u201313.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Rak, Yoel. 1983. <em>The <\/em><em>A<\/em><em>ustralopithecine <\/em><em>F<\/em><em>ace<\/em>. New York: Academic Press.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Rak, Yoel. 1988. \u201cOn Variation in the Masticatory System of <em>Australopithecus boisei<\/em>.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by M. Ruse and J. Travis, 193\u2013198<em>.<\/em> New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Semaw, Sileshi. 2000. \u201cThe World\u2019s Oldest Stone Artefacts from Gona, Ethiopia: Their Implications for Understanding Stone Technology and Patterns of Human Evolution between 2.6 Million Years Ago and 1.5 Million Years Ago.\u201d <em>Journal of Archaeological Science<\/em> 27(12): 1197\u20131214.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Shipman, Pat. 2002. <em>The Man Who Found the Missing Link: Eug<\/em><em>e<\/em><em>ne Dubois and <\/em><em>h<\/em><em>is Lifelong Quest to Prove Darwin Right<\/em>. New York: Simon &amp; Schuster.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Spoor, Fred. 2015. \u201cPalaeoanthropology: The Middle Pliocene Gets Crowded.\u201d<em> Nature<\/em> 521 (7553): 432\u2013433.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Strait, David S., Frederick E. Grine, and Marc A. Moniz. 1997. A Reappraisal of Early Hominid Phylogeny.\u201d <em>Journal of <\/em><em>H<\/em><em>uman <\/em><em>E<\/em><em>volution<\/em> 32 (1): 17\u201382.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Thackeray, J. Francis. 2000. \u201c\u2018Mrs. Ples\u2019 from Sterkfontein: Small Male or Large Female?\u201d <em>The South African Archaeological <\/em><em>Bulletin<\/em> 55: 155\u2013158.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Thackeray, J. Francis, Jos\u00e9 Braga, Jacques Treil, N. Niksch, and J. H. Labuschagne. 2002. \u201c\u2018Mrs. Ples\u2019 (Sts 5) from Sterkfontein: An Adolescent Male?\u201d <em>South African Journal of Science<\/em> 98 (1\u20132): 21\u201322.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Toth, Nicholas. 1985. \u201cThe Oldowan Reassessed.\u201d <em>Journal of Archaeological Science<\/em>\u00a012 (2): 101\u2013120.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Vrba, E. S. 1988. \u201cLate Pliocene Climatic Events and Hominid Evolution.\u201d In <em>The <\/em><em>E<\/em><em>volutionary <\/em><em>H<\/em><em>istory of the <\/em><em>R<\/em><em>obust Australopithecines<\/em>, edited by F. E. Grine, 405\u2013426. New York: Aldine.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Vrba, Elisabeth S. 1998. \u201cMultiphasic Growth Models and the Evolution of Prolonged Growth Exemplified by Human Brain Evolution.\u201d <em>Journal of Theoretical Biology<\/em> 190 (3): 227\u2013239.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Vrba, Elisabeth S. 2000. \u201cMajor Features of Neogene Mammalian Evolution in Africa.\u201d In <em>Cenozoic <\/em><em>G<\/em><em>eology of <\/em><em>S<\/em><em>outhern Africa<\/em>, edited by T. C. Partridge and R. Maud, 277\u2013304<em>.<\/em> Oxford: Oxford University Press.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Walker, Alan C., and Richard E. Leakey. 1988. \u201cThe Evolution of <em>Australopithecus boisei<\/em>.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by F. E. Grine, 247\u2013258. New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000\">Walker, Alan, Richard E. Leakey, John M. Harris, and Francis H. Brown. 1986. \u201c2.5-my <em>Australopithecus boisei<\/em> from West of Lake Turkana, Kenya.\u201d <em>Nature<\/em> 322 (6079): 517\u2013522.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Ward, Carol, Meave Leakey, and Alan Walker. 1999. \u201cThe New Hominid Species <em>Australopithecus anamensis<\/em>.\u201d <em>Evolutionary Anthropology<\/em> 7 (6): 197\u2013205.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">White, Tim D. 1988. \u201cThe Comparative Biology of \u2018Robust\u2019 Australopithecus: Clues from Content.\u201d In <em>Evolutionary History of the \u201cRobust\u201d Australopithecines<\/em>, edited by F. E. Grine, 449\u2013483. New York: Aldine de Gruyter.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">White, Tim D., Gen Suwa, and Berhane Asfaw. 1994. \u201c<em>Australopithecus ramidus<\/em>, a New Species of Early Hominid from Aramis, Ethiopia.\u201d <em>Nature<\/em> 371 (6495): 306\u2013312.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Wood, Bernard. 2010. \u201cReconstructing Human Evolution: Achievements, Challenges, and Opportunities.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 10 (2): 8902\u20138909.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Wood, Bernard, and Eve K. Boyle. 2016. \u201cHominin Taxic Diversity: Fact or Fantasy?\u201d <em>Yearbook of Physical Anthropology<\/em> 159 (S61): 37\u201378.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Wood, Bernard, and Kes Schroer. 2017. \u201cParanthropus: Where Do Things Stand?\u201d In <em>Human Paleontology and Prehistory<\/em>, edited by A. Marom and E. Hovers, 95\u2013107. New York: Springer, Cham.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000\">Acknowledgements<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">All of the authors in this section are students and early career researchers in paleoanthropology and related fields in South Africa (or at least have worked in South Africa). We wish to thank everyone who supports young and diverse talent in this field and would love to further acknowledge Black, African, and female academics who have helped pave the way for us.<\/span><\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_864\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_864\"><div tabindex=\"-1\"><p>Large animals such as mammoths and mastodons.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1774\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1774\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Bonnie Yoshida-Levine Ph.D., Grossmont College<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><em>This chapter is a revision from <\/em><em>\"<\/em><a class=\"rId7\" style=\"color: #000000;\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-15\/\"><em>Chapter 10: Early Members of the Genus Homo<\/em><\/a><em>\" by Bonnie Yoshida-Levine. In <\/em><a class=\"rId8\" style=\"color: #000000;\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, first edition<\/em><\/a><em>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under <\/em><a class=\"rId9\" style=\"color: #000000;\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\"><em>CC BY-NC 4.0<\/em><\/a><em>.<\/em><\/span><\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\"><span style=\"color: #ffffff;\">Learning Objectives<\/span><\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li>Describe how early Pleistocene climate change influenced the evolution of the genus Homo.<\/li>\n<li>Identify the characteristics that define the genus Homo.<\/li>\n<li>Describe the skeletal anatomy of Homo habilis\u00a0and Homo erectus based on the fossil evidence.<\/li>\n<li>Assess opposing points of view about how early Homo should be classified.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p><span style=\"color: #000000;\">The boy was no older than nine years when he perished by the swampy shores of the lake. After death, his slender, long-limbed body sank into the mud of the lake shallows. His bones fossilized and lay undisturbed for 1.5 million years. In the 1980s, fossil hunter Kamoya Kimeu, working on the western shore of Lake Turkana, Kenya, glimpsed a dark-colored piece of bone eroding in a hillside. This small skull fragment led to the discovery of what is arguably the world\u2019s most complete early hominin fossil\u2014a youth identified as a member of the species <em>Homo erectus<\/em>. Now known as Nariokotome Boy, after the nearby lake village, the skeleton has provided a wealth of information about the early evolution of our own genus, <em>Homo <\/em>(see Figure 10.1). Today, a stone monument with an inscription in three languages\u2014English, Swahili, and the local Turkana language\u2014marks the site of this momentous fossil discovery.<\/span><\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"aligncenter\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/06\/image8-1.jpg\" alt=\"Front view of near-complete skeleton\" width=\"407\" height=\"407\" \/><\/p>\n<figure style=\"width: 405px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image13-1.jpg\" alt=\"Reconstructed head and shoulders of a young Homo erectus.\" width=\"405\" height=\"308\" \/><figcaption class=\"wp-caption-text\">Figure 10.1a-b: a. Skeleton of a young male Homo erectus known as \u201cNariokotome Boy\u201d; b. an artist\u2019s depiction of how he may have looked during his life. This is the most complete hominin fossil from this time period ever found. Credit: a.<a href=\"https:\/\/humanorigins.si.edu\/evidence\/human-fossils\/fossils\/knm-wt-15000\"> KNM-WT 15000 Turkana Boy Skeleton<\/a> by<a href=\"https:\/\/www.si.edu\/\"> Smithsonian<\/a> [exhibit:<a href=\"https:\/\/humanorigins.si.edu\/research\"> Human Evolution<\/a> Evidence, Human Fossils, Fossils, KNM-WT 15000] is<a href=\"https:\/\/www.si.edu\/termsofuse\/\"> copyrighted and used for educational and non-commercial purposes as outlined by the Smithsonian<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Homo-erectus_Turkana-Boy_%28Ausschnitt%29_Fundort_Nariokotome,_Kenia,_Rekonstruktion_im_Neanderthal_Museum.jpg\">Homo-erectus Turkana-Boy (Ausschnitt) Fundort Nariokotome, Kenia, Rekonstruktion im Neanderthal Museum<\/a> by Neanderthal Museum is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Chapter 9 described our oldest human ancestors, primarily members of the genus <em>Australopithecus<\/em>, who lived between 2 million and 4 million years ago. This chapter introduces the earliest members of the genus <em>Homo<\/em>, focusing on <em>Homo habilis<\/em> and <em>Homo erectus<\/em>.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000;\">Defining the Genus <em>Homo<\/em><\/span><\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Because Anthropology is fundamentally concerned with what makes us human, defining our own genus takes on special significance for anthropologists. Ever since scientists acknowledged the existence of extinct species of humans, they have debated which of them display sufficient \u201chumanness\u201d to merit classification in the genus <em>Homo<\/em>. When grouping species into a common genus, biologists consider criteria such as physical characteristics (morphology), evidence of recent common ancestry, and adaptive strategy (use of the environment). However, there is disagreement about which of those criteria should be prioritized, as well as how specific fossils should be interpreted in light of the criteria.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Nevertheless, there is general agreement that species classified as <em>Homo<\/em> should share characteristics that are broadly similar within our species. These include the following:<\/span><\/p>\n<ul>\n<li class=\"import-Normal\" style=\"background-color: transparent; text-align: left; text-indent: 18pt;\"><span style=\"color: #000000;\">a relatively large brain size, <del>indicating a high degree of intelligence;<\/del><\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent; text-align: left; text-indent: 18pt;\"><span style=\"color: #000000;\">a smaller and flatter face<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent; text-align: left; text-indent: 18pt;\"><span style=\"color: #000000;\">smaller jaws and teeth<\/span><\/li>\n<li class=\"import-Normal\" style=\"background-color: transparent; text-align: left; text-indent: 18pt;\"><span style=\"color: #000000;\">increased reliance on culture, particularly the use of stone tools, to exploit a greater diversity of environments (adaptive zone).<\/span><\/li>\n<\/ul>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Some researchers would include larger overall body size and limb proportions (longer legs\/shorter arms) in this list. While these criteria seem relatively clear-cut, evaluating them in the fossil record has proved more difficult, particularly for the earliest members of the genus. There are several reasons for this. First, many fossil specimens dating to this time period are incomplete and poorly preserved. Second, early <em>Homo<\/em> fossils appear quite variable in brain size, facial features, and teeth and body size, and there is not yet consensus about how to best make sense of this diversity. Finally, there is growing evidence that the evolution of the genus <em>Homo<\/em> proceeded in a mosaic pattern: in other words, these characteristics did not appear all at once in a single species; rather, they were patchily distributed in different species from different regions and time periods. Consequently, different researchers have come up with conflicting classification schemes depending on which criteria they think are most important.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000; background-color: #ff99cc;\">In this chapter, we will take several pathways toward examining the origin and evolution of the genus <em>Homo<\/em>. First, we will explore the environmental conditions of the Pleistocene epoch in which the genus <em>Homo<\/em> evolved. Next we will examine the fossil evidence for the two principal species traditionally identified as early Homo: <em>Homo habilis<\/em> and <em>Homo erectus<\/em>. Then we will use data from fossils and archaeological sites to reconstruct the behavior of early members of <em>Homo<\/em>, including tool manufacture, subsistence practices, migratory patterns, and social structure. Finally, we will consider these together in an attempt to characterize the key adaptive strategies of early <em>Homo<\/em> and how they put our early ancestors on the trajectory that led to our own species, <em>Homo sapiens<\/em>.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000;\">Climate Change and Human Evolution<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">A key goal in the study of human origins is to learn about the environmental pressures that may have shaped human evolution. As indicated in Chapter 7, scientists use a variety of techniques to reconstruct ancient environments. These include stable isotopes, core samples from oceans and lakes, windblown dust, analysis of geological formations and volcanoes, and fossils of ancient plant and animal communities. Such studies have provided valuable information about the environmental context of early <em>Homo<\/em>.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">The early hominin species covered in Chapter 9, such as <em>Ardipithecus ramidus<\/em> and <em>Australopithecus afarensis<\/em>, evolved during the late <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1404\">Pliocene<\/a><\/strong> epoch. The Pliocene (5.3 million to 2.6 million years ago) was marked by cooler and drier conditions, with ice caps forming permanently at the poles. Still, Earth\u2019s climate during the Pliocene was considerably warmer and wetter than at present.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">The subsequent <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1549\">Pleistocene<\/a> <\/strong>epoch (2.6 million years to 11,000 years ago) ushered in major environmental change. The Pleistocene is popularly referred to as the Ice Age. Since the term \u201cIce Age\u201d tends to conjure up images of glaciers and woolly mammoths, one would naturally assume that this was a period of uniformly cold climate around the globe. But this is not actually the case. Instead, climate became much more variable, cycling abruptly between warm\/wet (interglacial) and cold\/dry (glacial) cycles. These patterns were influenced by changes in Earth\u2019s elliptical orbit around the sun. As is shown in Figure 10.2, each cycle averaged about 41,000 years during the early Pleistocene; the cycles then lengthened to about 100,000 years starting around 1.25 million years ago. Since mountain ranges, wind patterns, ocean currents, and volcanic activity can all influence climate patterns, there were wide-ranging regional and local effects.<\/span><\/p>\n<figure style=\"width: 655px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image7-3.png\" alt=\"Graph depicts five million years of climate change from sediment cores.\" width=\"655\" height=\"197\" \/><figcaption class=\"wp-caption-text\">Figure 10.2: Temperature estimates during the last five million years, extrapolated from deep-sea core data. Lower temperatures and increased temperature oscillations start at 2.6 million years ago. Glacial\/interglacial cycles during the early part of the epoch are shorter, each averaging about 41,000 years. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Five_Myr_Climate_Change.png\">Five Myr Climate Change<\/a> by<a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Dragons_flight\"> Dragons flight<\/a> (Robert A. Rohde), based on data from Lisiechi and Raymo (2005), is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\"> CC BY-SA 3.0 License.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Data on ancient geography and climate help us understand how our ancestors moved and migrated to different parts of the world\u2014as well as the constraints under which they operated. When periods of global cooling dominated, sea levels were lower as more water was captured as glacial ice. This exposed continental margins and opened pathways between land masses. During glacial periods, the large Indonesian islands of Sumatra, Java, and Borneo were connected to the Southeast Asian mainland, while New Guinea was part of the southern landmass of greater Australia. There was a land bridge connection between Britain and continental Europe, and an icy, treeless plain known as Beringia connected Northern Asia and Alaska. At the same time, glaciation made some northern areas inaccessible to human habitation. For example, there is evidence that hominin species were in Britain 950,000 years ago, but it does not appear that Britain was continuously occupied during this period. <span style=\"text-decoration: underline;\">(It is speculated)<\/span> These early humans may have died out or been forced to abandon the region during glacial periods.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">In Africa, paleoclimate research has determined that grasslands (shown in Figure 10.3) expanded and shrank multiple times during this period, even as they expanded over the long term (deMenocal 2014). From studies of fossils, paleontologists have been able to reconstruct Pleistocene animal communities and to consider how they were affected by the changing climate. Among the African animal populations, the number of grazing animal species such as antelope increased. Although the African and Eurasian continents are connected by land, the Sahara desert and the mountainous topography of North Africa serve as natural barriers to crossing. But the fossil record shows that at different times animal species have moved back and forth between Africa and Eurasia. During the early Pleistocene, there is evidence of African mammal species such as baboons, hippos, antelope, and African buffalo migrating out of Africa into Eurasia during periods of aridity (Belmaker 2010).<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 583px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image3-2.jpg\" alt=\"Dry grassy field with a few trees and mountains in the far distance.\" width=\"583\" height=\"406\" \/><figcaption class=\"wp-caption-text\">Figure 10.3: A savanna grassland in East Africa. Habitats such as this were becoming increasingly common during the Pleistocene. Credit: <a href=\"https:\/\/www.flickr.com\/photos\/ilri\/5130992564\">Savanna grasslands of East Africa<\/a> by<a href=\"https:\/\/www.flickr.com\/photos\/ilri\/\"> International Livestock Research Institute (ILRI)\/Elsworth<\/a> is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\"> CC BY-NC-SA 2.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">This changing environment was <del>undoubtedly<\/del> challenging for our ancestors, but it offered new opportunities to make a living. <span style=\"background-color: #ffff00;\">One solution adopted by some hominins was to specialize in feeding on the new types of plants growing in this landscape. The robust australopithecines (described in Chapter 9) likely developed their large molar teeth with thick enamel in order to exploit this particular dietary niche.<\/span><\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000; background-color: #ffff00;\">Members of the genus <em>Homo <\/em>took a different route. Faced with the unstable African climate and shifting landscape, they evolved bigger brains that enabled them to rely on cultural solutions such as crafting stone tools that opened up new foraging opportunities. This strategy of behavioral flexibility served them well during this unpredictable time and led to new innovations such as increased meat-eating, cooperative hunting, and the exploitation of new environments outside Africa. <\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000;\"><em>Homo habilis<\/em>: The Earliest Members of Our Genus<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><em>Homo habilis<\/em> has traditionally been considered the earliest species placed in the genus <em>Homo<\/em>. However, as we will see, there is substantial disagreement among paleoanthropologists about the fossils classified as <em>Homo habilis<\/em>, including whether they come from a single species or multiple, or even whether they should be part of the genus <em>Homo <\/em>at all.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><em>Homo habilis<\/em> has a somewhat larger brain size\u2014an average of 650 cubic centimeters (cc)\u2014compared to <em>Australopithecus<\/em> with less than 500 cc. Additionally, the skull is more rounded and the face less prognathic. However, the postcranial remains show a body size and proportions similar to <em>Australopithecus<\/em>.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Known dates for fossils identified as <em>Homo habilis<\/em> range from about 2.5 million years ago to 1.7 million years ago. Recently, a partial lower jaw dated to 2.8 million years from the site of Ledi-Gararu in Ethiopia has been tentatively identified as belonging to the genus <em>Homo<\/em> (Villmoare et al. 2015). If this classification holds up, it would push the origins of our genus back even further.<\/span><\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 554px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image16.jpg\" alt=\"Africa map with South Africa, Tanzania, Kenya, and Ethiopia shaded.\" width=\"554\" height=\"717\" \/><figcaption class=\"wp-caption-text\">Figure 10.4: Map showing major sites where <em>Homo habilis<\/em> fossils have been found. Ledi-Geraru is located in Ethiopia, Koobi Fora and Lake Turkana Basin are located in Kenya, the Olduvai Gorge is located in Tanzania, and Tuang, Malapa, Rising Star and Sterkfontein are located in South Africa. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-15\/\">Homo habilis site map (Figure 10.4)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Chelsea Barron at<a href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\"> GeoPlace, California State University, Chico<\/a> is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><span style=\"color: #000000;\"><strong><del>Discovery and Naming<\/del> <span style=\"text-decoration: underline;\">(just add paragraph not own section)<\/span><\/strong><\/span><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000; background-color: #ccffcc;\">The first fossils to be named <em>Homo habilis<\/em> were discovered at the site of Olduvai Gorge in Tanzania, East Africa, by members of a team led by Louis and Mary Leakey (Figure 10.4). The Leakey family had been conducting fieldwork in the area since the 1930s and had discovered other hominin fossils at the site, such as the robust <em>Paranthropos boisei<\/em>. The key specimen, a juvenile individual, was actually found by their 20-year-old son Jonathan Leakey. Louis Leakey invited South African paleoanthropologist Philip Tobias and British anatomist John Napier to reconstruct and analyze the remains. The fossil of the juvenile shown in Figure 10.5 (now known as OH-7) consisted of a lower jaw, parts of the parietal bones of the skull, and some hand and finger bones. The fossil was dated by potassium-argon dating to about 1.75 million years. In 1964, the team published their findings in the scientific journal <em>Nature <\/em>(Leakey et al. 1964)<em>. <\/em>As described in the publication, the new fossils had smaller molar teeth that were less \u201cbulgy\u201d than australopithecine teeth. Although the primary specimen was not yet fully grown, an estimate of its anticipated adult brain size would make it somewhat larger-brained than australopithecines such as <em>Austalopithecus africanus<\/em>. The hand bones were capable of a precision grip like a human\u2019s hand. This increased the likelihood that stone tools found earlier at Olduvai Gorge were made by this group of hominins. Based on these findings, the authors inferred that it was a new species that should be classified in the genus <em>Homo<\/em>. They gave it the name <em>Homo habilis<\/em>, meaning \u201chandy\u201d or \u201cskilled.\u201d<\/span><\/p>\n<p>&nbsp;<\/p>\n<\/div>\n<figure id=\"attachment_317\" aria-describedby=\"caption-attachment-317\" style=\"width: 641px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-317\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/10.5.jpg\" alt=\"Two lateral right views of skulls and a jaw with blackened teeth.\" width=\"641\" height=\"214\" \/><figcaption id=\"caption-attachment-317\" class=\"wp-caption-text\">Figure 10.5a-c: Homo habilis fossil specimens. From left to right they are: a. lateral right view of OH-24 (found at Olduvai Gorge), b. lateral right view of KNM-ER-1813 (from Koobi Fora, Kenya), and c. the jaw of OH-7, which was the type specimen found in 1960 at Olduvai Gorge, Tanzania. Credit: a. <a href=\"https:\/\/efossils.org\/page\/boneviewer\/Homo%20habilis\/OH%2024\">Homo habilis: OH 24 lateral right view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>. b. <a href=\"https:\/\/www.efossils.org\/page\/boneviewer\/Homo%20habilis\/KNM-ER%201813\">Homo habilis: KNM-ER 1813 lateral right view<\/a> by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>. c. <a href=\"https:\/\/boneclones.com\/product\/homo-habilis-oh-7-jaw-KO-196\">Homo habilis OH 7 Jaw<\/a> by<a href=\"https:\/\/boneclones.com\/\"> \u00a9BoneClones<\/a> is used by permission and available here under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<div class=\"__UNKNOWN__\">\n<h3 class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Controversies over Classification of <\/strong><strong><em>Homo habilis<\/em><\/strong><em><br style=\"clear: both;\" \/><\/em><\/span><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Since its initial discovery, many more Homo habilis were discovered in East and South African sites during the 1970s and 1980s (Figure 10.6). As more fossils joined the ranks of <em>Homo habilis<\/em>, several trends became apparent. First, the fossils were quite variable. While some resembled the fossil specimen first published by Leakey and colleagues, others had larger cranial capacity and tooth size. A well-preserved fossil skull from East Lake Turkana labeled KNM-ER-1470 displayed a larger cranial size along with a strikingly wide face. The diversity of the <em>Homo habilis<\/em> fossils prompted some scientists to question whether they displayed too much variation to all belong to the same species. They proposed splitting the fossils into at least two groups. The first group resembling the original small-brained specimen would retain the species name <em>Homo habilis<\/em>; the second group consisting of the larger-brained fossils such as KNM-ER-1470 would be assigned the new name of <em>Homo rudolfensis <\/em>(see Figure 10.7). Researchers who favored keeping all fossils in <em>Homo habilis<\/em> argued that sexual dimorphism, adaptation to local environments, or<strong> developmental plasticity<\/strong> could be the cause of the differences. For example, modern human body size and body proportions are influenced by variations in climates and nutritional circumstances.<\/span><\/p>\n<div style=\"text-align: left;\">\n<table class=\"aligncenter\" style=\"width: 467.5pt; height: 367px;\">\n<caption>Figure 10.6: Key Homo habilis fossil locations and the corresponding fossils and dates. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-15\/\">Homo habilis table (Figure 10.6)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Bonnie Yoshida-Levine is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>.<\/caption>\n<thead>\n<tr style=\"height: 0;\">\n<td class=\"Table1-C\" style=\"padding: 0pt 5.4pt; border: 0.5pt solid #000000; height: 60px; width: 130.2px;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Location of Fossils<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"padding: 0pt 5.4pt; border: 0.5pt solid #000000; height: 60px; width: 52.7px;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"padding: 0pt 5.4pt; border: 0.5pt solid #000000; height: 60px; width: 320.9px;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong><br \/>\nDescription <\/strong><\/span><\/p>\n<p>&nbsp;<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table1-R\" style=\"height: 0;\">\n<td class=\"Table1-C\" style=\"padding: 0pt 5.4pt; border: 0.5pt solid #000000; height: 61px; width: 130.2px;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Ledi-Gararu, Ethiopia<\/span><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"padding: 0pt 5.4pt; border: 0.5pt solid #000000; height: 61px; width: 52.7px;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">2.8 mya<\/span><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"padding: 0pt 5.4pt; border: 0.5pt solid #000000; height: 61px; width: 320.9px;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Partial lower jaw with evidence of both <em>Australopithecus<\/em> and <em>Homo<\/em> traits; tentatively considered oldest Early <em>Homo<\/em> fossil evidence.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0;\">\n<td class=\"Table1-C\" style=\"padding: 0pt 5.4pt; border: 0.5pt solid #000000; height: 77px; width: 130.2px;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Olduvai Gorge, Tanzania<\/span><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"padding: 0pt 5.4pt; border: 0.5pt solid #000000; height: 77px; width: 52.7px;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">1.7 mya to 1.8 mya<\/span><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"padding: 0pt 5.4pt; border: 0.5pt solid #000000; height: 77px; width: 320.9px;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Several different specimens classified as Homo habilis, including the type specimen found by Leakey, a relatively complete foot, and a skull with a cranial capacity of about 600 cc.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0;\">\n<td class=\"Table1-C\" style=\"padding: 0pt 5.4pt; border: 0.5pt solid #000000; height: 93px; width: 130.2px;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Koobi Fora, Lake Turkana Basin, Kenya<\/span><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"padding: 0pt 5.4pt; border: 0.5pt solid #000000; height: 93px; width: 52.7px;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">1.9 mya<\/span><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"padding: 0pt 5.4pt; border: 0.5pt solid #000000; height: 93px; width: 320.9px;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Several fossils from the Lake Turkana basin show considerable size differences, leading some anthropologists to classify the larger specimen (KNM-ER-1470) as a separate species,<em> Homo rudolfensis<\/em>.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0;\">\n<td class=\"Table1-C\" style=\"padding: 0pt 5.4pt; border: 0.5pt solid #000000; height: 61px; width: 130.2px;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Sterkfontein and other possible South African cave sites<\/span><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"padding: 0pt 5.4pt; border: 0.5pt solid #000000; height: 61px; width: 52.7px;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">about 1.7 mya<\/span><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"padding: 0pt 5.4pt; border: 0.5pt solid #000000; height: 61px; width: 320.9px;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">South African caves have yielded fragmentary remains identified as <em>Homo habilis<\/em>, but secure dates and specifics about the fossils are lacking.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr style=\"height: 15px;\">\n<td style=\"height: 15px; width: 132.133px;\"><\/td>\n<td style=\"height: 15px; width: 54.6333px;\"><\/td>\n<td style=\"height: 15px; width: 322.333px;\"><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<figure style=\"width: 228px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image10-1.jpg\" alt=\"Front view of black and white skull, missing lower jawbone.\" width=\"228\" height=\"228\" \/><figcaption class=\"wp-caption-text\">Figure 10.7: Cast of the Homo habilis cranium KNM-ER-1470. This cranium has a wide, flat face, larger brain size, and larger teeth than other Homo habilis fossils, leading some scientists to give it a separate species name, Homo rudolfensis. Credit: <a href=\"https:\/\/boneclones.com\/product\/homo-rudolfensis-skull-knm-er-1470-BH-013\">Homo rudolfensis Cranium KNM-ER 1470<\/a> by<a href=\"https:\/\/boneclones.com\/\"> \u00a9BoneClones<\/a> is used by permission and available here under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Given the incomplete and fragmentary fossil record from this time period, it is not surprising that classification has proved contentious. As a scholarly consensus has not yet emerged on the classification status of early <em>Homo<\/em>, this chapter makes use of the single (inclusive) <em>Homo habilis<\/em> species designation.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">There is also disagreement on whether <em>Homo habilis<\/em> legitimately belongs in the genus <em>Homo<\/em>. Most of the fossils first classified as <em>Homo habilis<\/em> were skulls and teeth. When arm, leg, and foot bones were later found, making it possible to estimate body size, the specimens turned out to be quite small in stature with long arms and short legs. Analysis of the relative strength of limb bones suggested that the species, though bipedal, was much more adapted to arboreal climbing than <em>Homo erectus<\/em> and <em>Homo sapiens <\/em>(Ruff 2009). This has prompted some scientists to assert that <em>Homo habilis<\/em> behaved more like an australopithecine\u2014with a shorter gait and the ability to move around in the trees (Wood and Collard 1999). They were also skeptical of the claim that the brain size of <em>Homo habilis<\/em> was much larger than that of <em>Australopithecus<\/em>. They have proposed reclassifying some or all of the <em>Homo habilis<\/em> fossils into the genus <em>Australopithecus<\/em>, or even placing them into a newly created genus (Wood 2014).<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Other scholars have interpreted the fossil evidence differently. A recent reanalysis of <em>Homo habilis\/rudolfensis<\/em> fossils concluded that they sort into the genus <em>Homo<\/em> rather than <em>Australopithecus <\/em>(see Hominin Species Summaries at chapter end). In particular, statistical analysis performed indicates that the <em>Homo habilis<\/em> fossils differ significantly in average cranial capacity from the australopithecines. They also note that some australopithecine species such as the recently discovered <em>Australopithecus sediba<\/em> have relatively long legs, so body size may not have been as significant as brain- and tooth-size differences (Anton et al. 2014).<\/span><\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\"><span style=\"color: #000000;\">Special Topic: Kamoya Kimeu<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Kamoya Kimeu (1938\u20132022) is arguably the most prolific fossil hunter in the history of paleoanthropology (Figure 10.8). In addition to his many decades of work as a field excavator and project supervisor in East Africa, he also trained field workers and scholars and has served as curator for prehistoric sites for the National Museum of Kenya.<\/span><\/p>\n<figure style=\"width: 228px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image4-1.jpg\" alt=\"Man smiling at camera with lake and mountain in the background.\" width=\"228\" height=\"351\" \/><figcaption class=\"wp-caption-text\">Figure 10.8: Kamoya Kimeu (1938-2022). Credit: Photograph of Kamoya Kimeu by \u00a9Dr. Mark Teaford is used by permission.<\/figcaption><\/figure>\n<p><span style=\"color: #000000;\">Kamoya Kimeu was born in 1938 in rural southeastern Kenya. Despite a formal education that did not go past the sixth grade, he had an aptitude for languages and familiarity with the plants and animals in the East African bush that led him to a job in Tanzania as a field excavator for Louis and Mary Leakey in 1960. In the years that followed, Kimeu found dozens of major hominin fossils. These included a <em>Paranthropus boisei <\/em>mandible at Olduvai Gorge, <em>Homo habilis <\/em>specimen KNM-ER-1813 from the Turkana Basin (shown in Figure 10.5), and a key early modern<em> Homo sapiens<\/em> fossil from the Omo Valley, Ethiopia. Kimeu\u2019s most famous fossil discovery was the skeleton of a young <em>Homo erectus<\/em> by the Nariokotome river bed in 1984. This finding was highly significant because it was a nearly complete early hominin skeleton and provided insight into child development within this species. In recognition of his work, Kimeu was awarded the National Geographic Society La Gorce Medal by U.S. President Ronald Reagan in 1985.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Traditionally, there has been a divide between African field workers and foreign research scientists, who would typically conduct seasonal field work in Africa, then travel back to their home institutions to publish their findings. Although Kimeu received widespread acclaim for the Nariokotome discovery, as well as a personal acknowledgement in the publication of the find in the journal <em>Nature<\/em>, he was not credited as an author. More recently, Kimeu\u2019s intellectual contributions to the field of paleoanthropology have been recognized. In 2021, he received an honorary doctorate degree from Case Western Reserve University in Ohio. Kimeu\u2019s most lasting legacy may be his mentorship of countless field workers and students. Today, there are a small but growing number of Black African paleoanthropologists taking on principal roles in the science of human origins.<\/span><\/p>\n<\/div>\n<h2 class=\"import-Normal\"><span style=\"color: #000000;\"><em>Homo habilis<\/em> Culture and Lifeways<br \/>\n<\/span><\/h2>\n<h3 class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Early Stone Tools<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\"><span style=\"background-color: #ffff00;\">The larger brains and smaller teeth of early <em>Homo <\/em>are linked to a different adaptive strategy than that of earlier hominins: one dependent on modifying rocks to make stone tools and exploit new food sources.<\/span> As discussed in Chapter 9, the 3.3-million-year-old stone tools from the Lomekwi 3 site in Kenya were made by earlier hominin species than <em>Homo<\/em>. However, stone tools become more frequent at sites dating to about 2 million years ago, the time of <em>Homo habilis <\/em>(Roche et al. 2009). This suggests that these hominins were increasingly reliant on stone tools to make a living.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Stone tools are assigned a good deal of importance in the study of human origins. Examining the form of the tools, the raw materials selected, and how they were made and used can provide insight into the thought processes of early humans and how they modified their environment in order to survive. Paleoanthropologists have traditionally classified collections of stone tools into industries, based on their form and mode of manufacture. There is not an exact correspondence between a tool industry and a hominin species; however, some general associations can be made between tool industries and particular hominins, locations, and time periods.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1394\">Oldowan<\/a><\/strong> tool industry is named after the site of Olduvai Gorge in Tanzania where the tools were first discovered. The time period of the Oldowan is generally estimated to be 2.5 mya to 1.6 mya. The tools of this industry are described as \u201cflake and chopper\u201d tools\u2014the choppers consisting of stone cobbles with a few flakes struck off them (Figure 10.9). To a casual observer, these tools might not look much different from randomly broken rocks. However, they are harder to make than their crude appearance suggests. The rock selected as the core must be struck by the rock serving as a hammerstone at just the right angle so that one or more flat flakes are removed. This requires selecting rocks that will fracture predictably instead of chunking, as well as the ability to plan ahead and envision the steps needed to create the finished product. The process leaves both the core and the flakes with sharp cutting edges that can be used for a variety of purposes.<\/span><\/p>\n<figure style=\"width: 505px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image11-1-1.png\" alt=\"Three stones with chunks missing from the tops and sides.\" width=\"505\" height=\"281\" \/><figcaption class=\"wp-caption-text\">Figure 10.9: Drawing of an Oldowan-style tool. This drawing shows a chopper; the flakes removed from the cores functioned as cutting tools. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Chopping_tool.gif\">Chopping tool<\/a> by Jos\u00e9-Manuel Benito \u00c1lvarez is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.5\/legalcode\">CC BY-SA 2.5 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Stone Tool Use and the Diet of Early <\/strong><strong><em>Homo<\/em><\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">What were the hominins doing with the tools? One key activity seems to have been butchering animals. Studies of animal bones at the site show cut marks on bones, and leg bones are often cracked open, suggesting that they were extracting the marrow from the bone cavities. It is interesting to consider whether the hominins hunted these animals or acquired them through other means. The butchered bones come from a variety of African mammals, ranging from small antelope to animals as big as wildebeest and elephants! It is difficult to envision slow, small-bodied <em>Homo habilis<\/em> with their Oldowan tools bringing down such large animals. One possibility is that the hominins were scavenging carcasses from lions and other large cats. Paleoanthropologist Robert Blumenschine has investigated this hypothesis by observing the behavior of present-day animal carnivores and scavengers on the African savanna. When lions abandon a kill after eating their fill, scavenging animals arrive almost immediately to pick apart the carcass. By the time slow-footed hominins arrived on the scene, the carcass would be mostly stripped of meat. However, if hominins could use stone tools to break into the leg bone cavities, they could get to the marrow, a fatty, calorie-dense source of protein (Blumenschine et al. 1987). Reconstructing activities that happened millions of years ago is obviously a difficult undertaking, and paleoanthropologists continue to debate whether scavenging or hunting was more commonly practiced during this time.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Regardless of how they were acquiring the meat, these activities suggest an important dietary shift from the way that the australopithecines were eating. The Oldowan toolmakers were exploiting a new ecological niche that provided them with more protein and calories. And it was not just limited to meat-eating\u2014stone tool use could have made available numerous other subsistence opportunities. A study of microscopic wear patterns on a sample of Oldowan tools indicates that they were used for processing plant materials such as wood, roots or tubers, and grass seeds and stems (Lemorini et al. 2014). In fact, it has been pointed out that the Oldowan toolmakers\u2019 cutting ability (whether for the purposes of consuming meat and plants or for making tools, shelters, or clothing) represents a new and unique innovation, never seen before in the natural world (Roche et al. 2009).<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000; background-color: #ff99cc;\">Overall, increasing the use of stone tools allowed hominins to expand their ecological niche and exert more control over their environment. As we\u2019ll see shortly, this pattern continued and became more pronounced with <em>Homo erectus<\/em>.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000;\"><em>Homo erectus<\/em>: Biological and Cultural Innovations<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Two million years ago, a new hominin appeared on the scene. Known as <em>Homo erectus<\/em>, the prevailing scientific view was that this species was much more like us. These hominins were equipped with bigger brains and large bodies with limb proportions similar to our own. Perhaps most importantly, their way of life is now one that is recognizably human, with more advanced tools, hunting, use of fire, and colonizing new environments outside of Africa.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">As will be apparent below, new data suggests that the story is not quite as simple. The fossil record for <em>Homo erectus<\/em> is much more abundant than that of <em>Homo habilis<\/em>, but it is also more complex and varied\u2014both with regard to the fossils as well as the geographic context in which they are found. <span style=\"background-color: #ff99cc;\">We will first summarize the anatomical characteristics that define <em>Homo erectus<\/em>, and then discuss the fossil evidence from Africa and the primary geographic regions outside Africa where the species has been located.<\/span><\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000;\"><strong><em>Homo erectus<\/em><\/strong><strong> Anatomy<\/strong><\/span><\/h3>\n<figure style=\"width: 289px\" class=\"wp-caption alignright\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image6-2.png\" alt=\"Lateral view of skull with large brow ridges.\" width=\"289\" height=\"289\" \/><figcaption class=\"wp-caption-text\">Figure 10.10: Replica of Homo erectus from Java, Indonesia. This cranium (known as Sangiran 17) dates to approximately 1.3 million to 1 million years ago. Note the large brow ridges and the occipital torus that gives the back of the skull a squared-off appearance. Credit: <a href=\"https:\/\/www.efossils.org\/page\/boneviewer\/Homo%20erectus\/Sangiran%2017\">Homo erectus: Sangiran 17 lateral left view<\/a>\u00a0 by \u00a9<a href=\"https:\/\/www.efossils.org\/\">eFossils<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/2.0\/\">CC BY-NC-SA 2.0 License<\/a> and is <a href=\"https:\/\/efossils.org\/page\/frequently-asked-questions\">used as outlined by eFossils<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Compared to <em>Homo habilis<\/em>, <em>Homo erectus<\/em> showed increased brain size, smaller teeth, and a larger body. However, it also displayed key differences from later hominin species including our own. Although the head of <em>Homo erectus<\/em> was less ape-like in appearance than the australopithecines, it did not resemble modern humans (Figure 10.10). Compared to <em>Homo habilis<\/em>, <em>Homo erectus<\/em> had a larger brain size: an average of about 900 cc compared to 650 cc to 750 cc. Instead of a rounded shape like our skulls, the <em>erectus <\/em>skull was long and low like a football, with a receding forehead, and a horizontal ridge called an <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1552\">occipital torus<\/a><\/strong> that gave the back of the skull a squared-off appearance. The cranial bones are thicker than those of modern humans, and some <em>Homo erectus<\/em> skulls have a slight thickening along the sagittal suture called a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1553\">sagittal keel<\/a><\/strong>. Large, shelf-like brow ridges hang over the eyes. The face shows less <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1554\"> prognathism<\/a><\/strong>, and the back teeth are smaller than those of <em>Homo habilis. <\/em>Instead of a pointed chin, like ours, the mandible of <em>Homo erectus<\/em> recedes back.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Apart from these features, there is significant variation among <em>Homo <\/em><em>erectus<\/em> fossils from different regions. Scientists have long noted differences between the fossils from Africa and those from Indonesia and China. For example, the Asian fossils tend to have a thicker skull and larger brow ridges than the African specimens, and the sagittal keel described above is more pronounced. <em>Homo erectus<\/em> fossils from the Republic of Georgia (described in the next section) also display distinctive characteristics. As with <em>Homo habilis<\/em>, this diversity has prompted a classification debate about whether or not <em>Homo erectus<\/em> should be split into multiple species. When African <em>Homo erectus<\/em> is characterized as a separate species, it is called <em>Homo ergaster<\/em>, while the Asian variant retains the <em>erectus <\/em>species name because it was discovered first. Here, the species name <em>Homo erectus<\/em> will be used for both variants.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\"><em>Homo erectus<\/em> was thought to have a body size and proportions more similar to modern humans. Unlike <em>Homo habilis<\/em> and the australopithecines, both of whom were small-statured with long arms and short legs, <em>Homo erectus<\/em> shows evidence of being fully committed to life on the ground. This meant long, powerfully muscled legs that enabled these hominins to cover more ground efficiently. Indeed, studies of the <em>Homo erectus<\/em> body form have linked several characteristics of the species to long-distance running in the more open savanna environment (Bramble and Lieberman 2004). Many experts think that hominins around this time had lost much of their body hair, were particularly efficient at sweating, and had darker-pigmented skin\u2014all traits that would support the active lifestyle of such a large-bodied hominin (see Special Topic box, \u201cHow We Became Sweaty, Hairless Primates\u201d).<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000;\">Special Topic: How We Became Hairless, Sweaty Primates <span style=\"text-decoration: underline;\">(include here)<\/span><\/span><\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Much of the information about the body form of <em>Homo erectus<\/em> comes from the Nariokotome fossil of the <em>Homo erectus<\/em> youth, described at the beginning of the chapter (see Figure 10.1). However, <em>Homo erectus<\/em> fossils are turning out to be more varied than previously thought. <em>Homo erectus <\/em>fossils from sites in Africa, as well as from Dmanisi, Georgia, show smaller body sizes than the Nariokotome boy. Even the Nariokotome skeleton itself has been reassessed: some now predict he would have been about 5 feet and 4 inches when fully grown rather than over 6 feet as initially hypothesized, although there is still disagreement about which measurement is more accurate. One explanation for the range of body sizes could be adaptation to a range of different local environments, just as humans today show reduced body size in poor nutritional environments (Anton and Snodgrass 2012).<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000;\"><strong><em>Homo erectus<\/em><\/strong><strong> in Africa <\/strong><\/span><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Although the earliest discoveries of <em>Homo erectus<\/em> fossils were from Asia, the greatest quantity and best-preserved fossils of the species come from East African sites. The earliest fossils in Africa identified as <em>Homo erectus <\/em>come from the East African site of Koobi Fora, around Lake Turkana in Kenya, and are dated to about 1.8 million years ago. Other fossil remains have been found in East African sites in Kenya, Tanzania, and Ethiopia. Other notable African <em>Homo erectus<\/em> finds are a female pelvis from the site of Gona, Ethiopia (Simpson et al. 2008), and a cranium with massive brow ridges from Olduvai Gorge known as Olduvai 9, thought to be about 1.4 million years old.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Until recently, <em>Homo erectus<\/em>\u2019 presence in southern Africa has not been well documented. However, work at the Drimolen cave site in South Africa has yielded new fossils of <em>Paranthropus robustus<\/em>, and the cranium of a 2\u20133 year old child tentatively identified as <em>Homo erectus<\/em>, dated to about 2 million years (Herries et al. 2020). If substantiated, this would be the oldest discovery to date of <em>Homo erectus<\/em> anywhere.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Regional Discoveries Outside Africa<\/strong><\/span><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">It is generally agreed that<em> Homo erectus<\/em> was the first hominin to migrate out of Africa and colonize Asia and later Europe (although recent discoveries in Asia may challenge this view). Key locations and discoveries of <em>Homo erectus<\/em> fossils, along with the fossils\u2019 estimated ages, are summarized in Figures 10.11 and 10.12.<\/span><\/p>\n<figure style=\"width: 594px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image17-3.jpg\" alt=\"World map with England, Spain, Georgia, Kenya, China, and Java shaded.\" width=\"594\" height=\"459\" \/><figcaption class=\"wp-caption-text\">Figure 10.11:\u00a0 Map showing the locations of Homo erectus fossils around Africa and Eurasia. <a href=\"https:\/\/docs.google.com\/document\/d\/1VUDKMBJYS_jNONjLxT04jQN0_z9Ua50BRN6auGSHUuU\/edit\">A full text description of this image is available<\/a>. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-15\/\">Homo erectus site map<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Chelsea Barron at<a href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\"> GeoPlace, California State University, Chico<\/a> is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<div style=\"text-align: left;\">\n<table class=\"aligncenter\" style=\"width: 467.5pt;\">\n<caption>Figure 10.12: Regional comparisons of Homo erectus fossils. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-15\/\">Regional comparisons of Homo erectus fossils (Figure 10.12)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/caption>\n<thead>\n<tr style=\"height: 0;\">\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Region<\/strong><\/span><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Sites<\/strong><\/span><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Dates<\/strong><\/span><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Significance of Fossils<\/strong><\/span><\/p>\n<p>&nbsp;<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table2-R\" style=\"height: 0;\">\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">East Africa<\/span><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">East and West Lake Turkana, Kenya; Olduvai Gorge, Tanzania<\/span><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">1.8 to 1.4 mya<\/span><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Earliest evidence of <em>H. erectus<\/em>; significant variation in skull and facial features.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 0;\">\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">South Africa<\/span><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Drimolen Cave, South Africa<\/span><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">2 mya<\/span><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Recent find of a 2\u20133 year old child would be oldest <em>H. erectus<\/em> anywhere to date.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 0;\">\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Western<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Eurasia<\/span><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Dmanisi, Republic of Georgia<\/span><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">1.75 mya<\/span><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Smaller brains and bodies than <em>H. erectus<\/em> from other regions.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 0;\">\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Western Europe<\/span><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Atapuerca, Spain (Sima del Elefante and Gran Dolina caves)<\/span><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">1.2 mya\u2013 400,000 ya<\/span><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Partial jaw from Atapuerca is oldest evidence of <em>H. erectus<\/em> in Western Europe.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Fossils from Gran Dolina (dated to about 800,000 years) sometimes referred to as <em>H. antecessor.<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 0;\">\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Indonesia<\/span><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Ngandong, Java; Sangiran, Java<\/span><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">1.6 mya<\/span><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Early dispersal of <em>H. erectus <\/em>to East Asia; Asian <em>H. erectus <\/em>features.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 0;\">\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">China<\/span><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Zhoukoudian, China;<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Loess Plateau (Lantian)<\/span><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">780,000\u2013 400,000 ya;<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">2.1 mya<\/span><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"padding: 0pt 5.4pt 0pt 5.4pt; border: solid #000000 0.5pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Large sample of <em>H. erectus<\/em> fossils and artifacts.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Recent evidence of stone tools from Loess Plateau suggests great antiquity of <em>Homo<\/em> in East Asia.<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<h4 class=\"import-Normal\"><span style=\"color: #000000;\"><em>Indonesia<\/em><\/span><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">The first discovery of<em> Homo erectus<\/em> was in the late 1800s in Java, Indonesia. A Dutch anatomist named Eugene Dubois searched for human fossils with the belief that since orangutans lived there, it might be a good place to look for remains of early humans. He discovered a portion of a skull, a femur, and other bone fragments on a riverbank. While the femur looked human, the top of the skull was smaller and thicker than that of a modern person. Dubois named the fossil <em>Pithecanthropus erectus<\/em> (\u201cupright ape-man\u201d), popularized in the media at the time as \u201cJava Man.\u201d After later discoveries of similar fossils in China and Africa, they were combined into a single species (retaining the <em>erectus<\/em> name) under the genus <em>Homo<\/em>.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Although <em>Homo erectus<\/em> has a long history in Indonesia, the region\u2019s geology has complicated the dating of fossils and sites. Fossils from the Sangiran Dome, Java, had previously been estimated to be as old as 1.8 million years, but scientists using new dating methods have arrived at a later date of about 1.3 mya (Matsu\u2019ura et al. 2020). On the recent end of the timeline, a cache of <em>H. erectus<\/em> fossils from the site of Ngandong in Java has yielded a surprisingly young date of 43,000 years, although a newer study with different dating methods concluded that they were between 117,000 to 108,000 years old (Rizal et al. 2020).<\/span><\/p>\n<h4 class=\"import-Normal\"><span style=\"color: #000000;\"><em>China<\/em><\/span><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">There is evidence of<em> Homo erectus <\/em>in China from several regions and time periods.<em> Homo erectus<\/em> fossils from northern China, collectively known as \u201cPeking Man,\u201d are some of the most famous human fossils in the world. Dated to about 400,000\u2013700,000 years ago, they were excavated from the site of Zhoukoudian, near the outskirts of Beijing. Hundreds of bones and teeth, including six nearly complete skulls, were excavated from a cave in the 1920s and 1930s. Much of the fossils\u2019 fame comes from the fact that they disappeared under mysterious circumstances. As Japan advanced into China during World War II, Chinese authorities, concerned for the security of the fossils, packed up the boxes and arranged for them to be transported to the United States. But in the chaos of the war, they vanished and were never heard about again. Fortunately, an anatomist named Frans Weidenreich had previously studied the bones and made casts and measurements of the skulls, so this valuable information was not lost. More recent excavations at Longgushan \u201cDragon Bone Cave\u201d at Zhoukoudian\u2014of tools, living sites, and food remains\u2014have revealed much about the lifestyle of <em>Homo erectus <\/em>during this time.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Despite this long history of research, China, compared to Africa, was perceived as somewhat peripheral to the study of hominin evolution. Although <em>Homo erectus<\/em> fossils have been found at several sites in China, with dates that make them comparable to those of Indonesian <em>Homo erectus<\/em>, none seemed to approximate the antiquity of African sites. The notable finds at sites like Nariokotome and Olorgesaille took center stage during the 1970s and 1980s, as scientists focused on elucidating the species\u2019 anatomy and adaptations in its African homeland. In contrast, fewer research projects were focused on East Asian sites (Dennell and Roebroeks 2005; Qiu 2016).<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">However, isolated claims of very ancient hominin occupation kept cropping up from different locations in Asia. While some were dismissed because of problems with dating methods or stratigraphic context, the 2018 publication of the discovery of 2.1-million-year-old stone tools from China caught everyone\u2019s attention. Based on paleomagnetic techniques that date the associated soils and windblown dust, these tools indicate that hominins in Asia predated those from the Georgian site of Dmanisi by at least 300,000 years (Zhu et al. 2018). In fact, the tools are older than any <em>Homo erectus<\/em> fossils anywhere. Since no fossils were found with the tools, it isn\u2019t known which species made them, but it opens up the intriguing possibility that hominins could have migrated out of Africa earlier than <em>Homo erectus<\/em>. These new discoveries are shaking up previously held views of the East Asian human fossil record.<\/span><\/p>\n<h4 class=\"import-Normal\"><span style=\"color: #000000;\"><em>Western Eurasia<\/em><\/span><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">An extraordinary collection of fossils from the site of Dmanisi in the Republic of Georgia has revealed the presence of <em>Homo erectus<\/em> in Western Eurasia between 1.75 million and 1.86 million years ago. Dmanisi is located in the Caucasus mountains in Georgia. When archaeologists began excavating a medieval settlement near the town in the 1980s and came across the bones of extinct animals, they shifted their focus from the historic to the prehistoric era, but they probably did not anticipate going back quite so far in time. The first hominin fossils were discovered in the early 1990s, and since that time, at least five relatively well-preserved crania have been excavated.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">There are several surprising things about the Dmanisi fossils. Compared to African <em>Homo erectus,<\/em> they have smaller brains and bodies. However, despite the small brain size, they show clear signs of <em>Homo erectus<\/em> traits such as heavy brow ridges and reduced facial prognathism. Paleoanthropologists have pointed to some aspects of their anatomy (such as the shoulders) that appear rather primitive, although their body proportions seem fully committed to terrestrial bipedalism. One explanation for these differences could be that the Dmanisi hominins represent a very early form of <em>Homo erectus<\/em> that left Africa before increases in brain and body size evolved in the African population.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Second, although the fossils at this location are from the same geological context, they show a great deal of variation in brain size and in facial features. One skull (Skull 5) has a cranial capacity of only 550 cc, smaller than many <em>Homo habilis <\/em>fossils, along with larger teeth and a protruding face. Scientists disagree on what these differences mean. Some contend that the Dmanisi fossils cannot all belong to a single species because each one is so different. Others assert that the variability of the Dmanisi fossils proves that they, along with all early Homo fossils, including <em>H. habilis<\/em> and <em>H.<\/em><em>rudolfensis, <\/em>could <em>all <\/em>be grouped into <em>Homo erectus<\/em> (Lordkipanidze et al. 2013). Regardless of which point of view ends up dominating, the Dmanisi hominins are clearly central to the question of how to define the early members of the genus <em>Homo<\/em>.<\/span><\/p>\n<h4 class=\"import-Normal\"><span style=\"color: #000000;\"><em>Europe<\/em><\/span><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Until recently, there was scant evidence of any <em>Homo erectus<\/em> presence in Europe, and it was assumed that hominins did not colonize Europe until much later than East Asia or Eurasia. One explanation for this was that the harsh climate of Western Europe served as a barrier to settlement. However, recent fossil finds from Spain suggest that <em>Homo erectus<\/em> could have made it into Europe over a million years ago. In 2008 a mandible from the Atapuerca region in Spain was discovered, dating to about 1.2 million years ago. A more extensive assemblage of fossils from the site of Gran Dolina in Atapuerca have been dated to about 800,000 years ago. In England in 2013 fossilized hominin footprints of adults and children dated to 950,000 years ago were found at the site of Happisburgh, Norfolk, which would make them the oldest human footprints found outside Africa (Ashton et al. 2014).<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">At this time, researchers aren\u2019t in agreement as to whether the first Europeans belonged to <em>Homo erectus<\/em> proper or to a later descendent species. Some scientists refer to the early fossils from Spain by the species name <em>Homo antecessor<\/em>.<\/span><\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\"><span style=\"color: #000000;\">Special Topic: How We Became Hairless, Sweaty Primates<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">As an anthropology instructor teaching human evolution, my students often ask me about human body hair: When did our ancestors lose it and why? It is assumed that our earliest ancestors were as hairy as modern-day apes. Yet, today, we lack thick hair on most parts of our bodies except in the armpits, pubic regions, and tops of our heads. Humans actually have about the same number of hair follicles per unit of skin as chimpanzees, but, the hairs on most of our body are so thin as to be practically invisible. When did we develop this peculiar pattern of hairlessness? Which selective pressures in our ancestral environment were responsible for this unusual characteristic?<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Many experts believe that the driving force behind our loss of body hair was the need to effectively cool ourselves. Along with the lack of hair, humans are also distinguished by being exceptionally sweaty: we sweat larger quantities and more efficiently than any other primate. Humans have a larger amount of eccrine sweat glands than other primates and these glands generate an enormous volume of watery sweat. Sweating produces liquid on the skin that cools the body off as it evaporates. It seems likely that hairlessness and sweating evolved together, as a recent DNA analysis has identified a shared genetic pathway between hair follicles and eccrine sweat gland production (Kamberov et al. 2015).<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Which particular environmental conditions led to such adaptations? In this chapter, we learned that the climate was a driving force behind many changes seen in the hominin lineage during the Pleistocene. At that time, the climate was increasingly arid and the forest canopy in parts of Africa was being replaced with a more open grassland environment, resulting in increased sun exposure for our ancestors. Compared to the earlier australopithecines, members of the genus <em>Homo<\/em> were also developing larger bodies and brains, starting to obtain meat by hunting or scavenging carcasses, and crafting sophisticated stone tools.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">According to Nina Jablonski, an expert on the evolution of human skin, the loss of body hair and increased sweating capacity are part of the package of traits characterizing the genus <em>Homo<\/em>. While larger brains and long-legged bodies made it possible for humans to cover long distances while foraging, this new body form had to cool itself effectively to handle a more active lifestyle. Preventing the brain from overheating was especially critical. The ability to keep cool may have also enabled hominins to forage during the hottest part of the day, giving them an advantage over savanna predators, like lions, that typically rest during this time (Jablonski 2010).<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">When did these changes occur? Although hair and soft tissue do not typically fossilize, several indirect methods have been used to explore this question. One method tracks a human skin color gene. Since chimpanzees have light skin under their hair, it is probable that early hominins also had light skin color. Apes and other mammals with thick fur coats have protection against the sun\u2019s rays. As our ancestors lost their fur, it is likely that increased melanin pigmentation was selected for as a way to shield our ancestors from harmful ultraviolet radiation. A recent genetic analysis determined that one of the genes responsible for melanin production originated about 1.2 million years ago (Rogers et al 2004).<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Another line of evidence tracks the coevolution of a rather unpleasant human companion\u2014the louse. A genetic study identified human body louse as the youngest of the three varieties of lice that infest humans, splitting off as a distinct variety around 70,000 years ago (Kittler et al. 2003). Because human body lice can only spread through clothing, this may have been about the time when humans started to regularly wear clothing. However, the split between human head and pubic lice is estimated to have occurred much earlier, about three million years ago (Bower 2003; Reed et al. 2007). When humans lost much of their body hair, lice that used to roam freely around the body were now confined to two areas: the head and pubic region. As a result of this separation, the lice population split into two distinct groups.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Other explanations have been suggested for the loss of human body hair. For example, being hairless makes it more difficult for skin parasites like lice, fleas, and ticks to live on us. Additionally, after bipedality evolved, hairless bodies would also make reproductive organs and female breasts more visible, suggesting that sexual selection may have played a role.<\/span><\/p>\n<\/div>\n<\/div>\n<h2 class=\"__UNKNOWN__\"><span style=\"color: #000000;\"><em>Homo erectus <\/em>Lifeways<br \/>\n<\/span><\/h2>\n<div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\"><span style=\"color: #000000; background-color: #ff99cc;\">Now, our examination of <em>Homo erectus<\/em> will turn to its lifeways\u2014how the species utilized its environment in order to survive. This includes making inferences about diet, technology, life history, environments occupied, and perhaps even social organization. As will be apparent, <em>Homo erectus <\/em>shows significant cultural innovations in these areas, some that you will probably recognize as more \u201chuman-like\u201d than any of the hominins previously covered.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Tool Technology: Acheulean Tool Industry<\/strong><\/span><\/h3>\n<figure style=\"width: 423px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image9-3.png\" alt=\"Front, back, side, and top views of oval-shaped stone core with chunks removed.\" width=\"423\" height=\"341\" \/><figcaption class=\"wp-caption-text\">Figure 10.13: Drawing of an Acheulean handaxe. This specimen is from Spain. When drawing a stone tool, artists typically show front and back faces, as well as top and side profiles. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Hand_axe_spanish.gif\">Hand axe spanish<\/a> by<a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Locutus_Borg\"> Jos\u00e9-Manuel Benito (user: Locutus Borg<\/a>) has been designated to the<a href=\"https:\/\/creativecommons.org\/share-your-work\/public-domain\/cc0\/\"> public domain (CC0)<\/a>.<\/figcaption><\/figure>\n<p>In early African sites associated with <em>Homo erectus<\/em>, stone tools such as flakes and choppers identified to the Oldowan Industry dominate. Starting at about 1.5 million years ago, some <em>Homo erectus<\/em> populations began making different forms of tools. These tools\u2014classified together as constituting the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1556\">Acheulean <\/a><\/strong> tool industry\u2014are more complex in form and more consistent in their manufacture. Unlike the Oldowan tools, which were cobbles modified by striking off a few flakes, Acheulean toolmakers carefully shaped both sides of the tool. This type of technique, known as bifacial flaking, requires more planning and skill on the part of the toolmaker; he or she would need to be aware of principles of symmetry when crafting the tool. One of the most common tool forms, the handaxe, is shown in Figure 10.13. As with the tool illustrated below, handaxes tend to be thicker at the base and then come to a rounded point at the tip. Besides handaxes, forms such as scrapers, cleavers, and flake tools are present at <em>Homo erectus<\/em> sites.<\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">One striking aspect of Acheulean tools is their uniformity. They are more standardized in form and mode of manufacture than the earlier Oldowan tools. For example, the aforementioned handaxes vary in size, but they are remarkably consistent in regard to their shape and proportions. They were also an incredibly stable tool form over time\u2014lasting well over a million years with little change.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Curiously, the Acheulean tools so prominent at African sites are mostly absent in <em>Homo erectus<\/em> sites in East Asia. Instead, Oldowan-type choppers and scrapers are found at those sites. If this technology seemed to be so important to African <em>Homo erectus<\/em>, why didn\u2019t East Asian <em>Homo erectus<\/em> also use the tools? One reason could be environmental differences between the two regions. It has been suggested that <em>Asian Homo<\/em> <em>erectus<\/em> populations used perishable material such as bamboo to make tools. Another possibility is that <em>Homo erectus<\/em> (or even an earlier hominin) migrated to East Asia before the Acheulean technology developed in Africa. The recent discovery of the 2.1-million-year-old tools in China gives credence to this last explanation.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">What (if anything) do the Acheulean tools tell us about the mind of <em>Homo erectus<\/em>? Clearly, they took a fair amount of skill to manufacture. Apart from the actual shaping of the tool, other decisions made by toolmakers can reveal their use of foresight and planning. Did they just pick the most convenient rocks to make their tools, or did they search out a particular raw material that would be ideal for a particular tool? Analysis of Acheulean stone tools suggest that at some sites, the toolmakers selected their raw materials carefully\u2014traveling to particular rock outcrops to quarry stones and perhaps even removing large slabs of rock at the quarries to get at the most desirable material. Such complex activities would require advanced planning and communication with other individuals. However, other <em>Homo erectus<\/em> sites lack evidence of such selectivity; instead of traveling even a short distance for better raw material, the hominins tended to use what was available in their immediate area (Shipton et al. 2018).<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">In contrast to <em>Homo erectus<\/em> tools, the tools of early modern <em>Homo sapiens<\/em> during the Upper Paleolithic display tremendous diversity across regions and time periods. Additionally, Upper Paleolithic tools and artifacts communicate information such as status and group membership. Such innovation and social signaling seem to have been absent in <em>Homo erectus<\/em>, suggesting that they had a different relationship with their tools than did <em>Homo sapiens<\/em> (Coolidge and Wynn 2017). Some scientists assert that these contrasts in tool form and manufacture may signify key cognitive differences between the species, such as the ability to use a complex language.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Subsistence and Diet<\/strong><\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">In reconstructing the diet of <em>Homo erectus<\/em>, researchers can draw from multiple lines of evidence. These include stone tools used by <em>Homo erectus<\/em>, animal bones and occasionally plant remains from <em>Homo erectus<\/em> sites, and the bones and teeth of the fossils themselves. These data sources suggest that compared to the australopithecines, <em>Homo erectus<\/em> consumed more animal protein. Coinciding with the appearance of <em>Homo erectus<\/em> fossils in Africa are archaeological sites with much more abundant stone tools and larger concentrations of butchered animal bones.<\/span><\/p>\n<figure style=\"width: 253px\" class=\"wp-caption alignright\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image5-2.png\" alt=\"Five men excavating and note-taking at an archaeological site.\" width=\"253\" height=\"380\" \/><figcaption class=\"wp-caption-text\">Figure 10.14: Excavations at the site of Olorgesailie, Kenya. Dated from between 1.2 million years ago and 490,000 years ago, Olorgesailie has some of the most abundant and well-preserved evidence of Homo erectus activity in the world. Fossils of large mammals, such as elephants, along with thousands of Acheulean tools, have been uncovered over the decades. Credit: <a href=\"https:\/\/humanorigins.si.edu\/research\/olorgesailie-kenya\">Elephant Butchery Site Olorgesailie, Kenya<\/a> by<a href=\"https:\/\/www.si.edu\/\"> Smithsonian<\/a> [exhibit:<a href=\"https:\/\/humanorigins.si.edu\/research\"> Human Evolution Research<\/a>,<a href=\"https:\/\/humanorigins.si.edu\/research\/east-african-research-projects\"> East African Research Projects<\/a>, Olorgesailie, Kenya] is<a href=\"https:\/\/www.si.edu\/termsofuse\/\"> copyrighted and used for educational and non-commercial purposes as outlined by the Smithsonian<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">It makes sense that a larger body and brain would be correlated with a dietary shift to more calorically dense foods. This is because the brain is a very energetically greedy organ. Indeed, our own human brains require more than 20% of one\u2019s calorie total intake to maintain. When biologists consider the evolution of intelligence in any animal species, it is often framed as a cost\/benefit analysis: For large brains to evolve, there has to be a compelling benefit to having them and a way to generate enough energy to fuel them.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">One solution that would allow for an increase in human brain size would be a corresponding reduction in the size of the digestive tract (gut). According to the \u201cexpensive tissue hypothesis,\u201d initially formulated by Leslie Aiello and Peter Wheeler (1995), a smaller gut would allow for a larger brain without the need for a corresponding increase in the organism\u2019s metabolic rate. More meat in the diet could also fuel the larger brain and body size seen in the genus <em>Homo<\/em>. Some researchers also believe that body fat percentages increased in hominins (particularly females) around this time, which would have allowed them to be better buffered against environmental disruption such as food shortages (Anton and Snodgrass 2012).<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">As indicated above, evidence from archaeology and the inferences about <em>Homo erectus<\/em> body size suggest increased meat eating. How much hunting did <em>Homo erectus<\/em> engage in compared to the earlier Oldowan toolmakers? Although experts continue to debate the relative importance of hunting versus scavenging, there seems to be stronger evidence of hunting for these hominins. For example, at sites such as Olorgesailie in Kenya (Figure 10.14), there are numerous associations of Acheulean tools with butchered remains of large animals.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">However, <em>Homo erectus<\/em> certainly ate more than just meat. Studies of the tooth surfaces and microscopic wear patterns on hominin teeth indicate that these hominins ate a variety of foods, including some hard, brittle plant foods (Unger and Scott 2009). This would make sense, considering the environment was changing to be more dominated by grasslands in some areas. Roots, bulbs, and tubers (known as underground storage organs) of open savanna plants may have been a primary food source. Indeed, hunter-gatherer groups such as the Hadza of Tanzania rely heavily on such foods, especially during periods when game is scarce. In the unstable environment of the early Pleistocene, dietary versatility would be a definite advantage.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Tool Use, Cooking, and Fire<\/strong><\/span><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">One key characteristic of the genus <em>Homo<\/em> is smaller teeth compared to <em>Australopithecus<\/em>. Why would teeth get smaller? In addition to new types of foods, changes in how food was prepared and consumed likely led to a decrease in tooth size. Think about how you would eat if you didn\u2019t have access to cutting tools. <span style=\"background-color: #ffff00;\">What you couldn\u2019t rip apart with your hands would have to be bitten off with your teeth\u2014actions that would require bigger, more powerful teeth and jaws. As stone tools became increasingly important, hominins began to cut up, tenderize, and process meat and plants, such that they did not have to use their teeth so vigorously.<\/span><\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Cooking food could also have contributed to the reduction in tooth and jaw size. In fact, anthropologist Richard Wrangham (2009) asserts that cooking played a crucial role in human evolution. Cooking provides a head start in the digestive process because of how heat begins to break down food before food even enters the body, and it can help the body extract more nutrients out of meat and plant foods such as starchy tubers.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Obviously cooking requires fire, and the earliest use of fire is a fascinating topic in the study of human evolution. Fire is not only produced by humans; it occurs naturally as a result of lightning strikes. Like other wild animals, early hominins must have been terrified of wildfires, but at some point in time they learned to control fire and put it to good use. Documenting the earliest evidence of fire has been a contentious issue in archaeology because of the difficulty in distinguishing between human-controlled fire and natural burning at hominin sites. Burned areas and ash deposits must have direct associations with human activity to make a case for deliberate fire use. Unfortunately, such evidence is rare at ancient hominin sites, which have been profoundly altered by humans, animals, and geological forces over millions of years. Recently, newer methods\u2014including microscopic analysis of burned rock and bone\u2014have revealed clear evidence of fire use at Koobi Fora, Kenya, dating to 1.5 million years ago (Hlubik et al. 2017).<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Migration out of Africa<\/strong><\/span><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><em>Homo erectus<\/em> is generally thought to be the first hominin species to have left Africa. It is hypothesized that they settled in places in Eurasia, such as the Republic of Georgia, Indonesia, and northern China, where fossil evidence of <em>Homo erectus<\/em> exists. But why would this species have traveled such vast distances to these far-flung regions? To answer this question, we have to consider what we have learned about the biology, culture, and environmental circumstances of <em>Homo erectus. <\/em><span style=\"background-color: #ffff00;\">The larger brain and body size of <em>Homo erectus<\/em> were fueled by a diet consisting of more meat, and their longer, more powerful legs made it possible to walk and run longer distances to acquire food. Since they were eating higher on the food chain, it was necessary for them to extend their home range to find sufficient game. Cultural developments\u2014including better stone tools and new technology such as fire\u2014gave them greater flexibility in adapting to different environments.<\/span> Finally, the major Pleistocene climate shift discussed earlier in the chapter certainly played a role. Changes in air temperature, precipitation, access to water sources, and other habitat alteration had far-reaching effects on animal and plant communities; this included <em>Homo erectus<\/em>. If hominins were relying more on hunting, the migration patterns of their prey could have led them to traverse increasingly long distances.<\/span><\/p>\n<h3 class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Life History<\/strong><\/span><\/h3>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1558\">life history <\/a><\/strong> of a species refers to its overall pattern of growth, development, and reproduction during its lifetime, with the assumption that these characteristics have been shaped by natural selection. The field of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1559\">human behavioral ecology<\/a><\/strong>, explored in more detail in Appendix C, examines the roots of human behavior and life history. Our species, <em>Homo sapiens<\/em>, is characterized by a unique life history pattern of slow development, an extended period of juvenile dependence, and a long lifespan. Whereas the offspring of great apes achieve self-sufficiency early, human children are dependent on their parents long after weaning. Additionally, human fathers and grandparents (particularly postmenopausal grandmothers) devote substantial time and energy to caring for their children.<\/span><\/p>\n<figure style=\"width: 310px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image15-3.png\" alt=\"One man is shooting a bow and arrow; another man is carrying a bow with a dog beside him.\" width=\"310\" height=\"465\" \/><figcaption class=\"wp-caption-text\">Figure 10.15: Hadza men practice bowing. Native to Tanzania, the Hadza have retained many traditional foraging practices. Although most do not subsist entirely upon wild foods today, their way of life may shed light on how humans lived for most of their evolutionary history. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Hadzabe1.jpg\">Hadzabe1<\/a> by<a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Idobi\"> Idobi<\/a> is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\"> CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Human behavioral ecologists who study modern hunter-gatherer societies have observed that foraging is no easy business (Figure 10.15). Members of these groups engage in complex foraging techniques that take many years to master. An extended juvenile period gives children the time to acquire these skills. It also allows time for large human brains to grow and mature. On the back end, a longer developmental period results in skilled, successful adults, capable of living a long time (Hill and Kaplan 1999). Despite the time and energy demands, females could have offspring at more closely spaced intervals if they could depend on help from fathers and grandmothers (Hawkes et al. 1998).<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">What can the study of <em>Homo erectus<\/em> reveal about its life history pattern? Well-preserved fossils such as the Nariokotome boy can provide some insights. We know that apes such as chimpanzees reach maturity more quickly than humans, and there is some evidence that the australopithecines had a growth rate more akin to that of chimpanzees. Scientists have conducted extensive studies of the Nariokotome skeleton\u2019s bones and teeth to assess growth and development. On the one hand, examination of the long bone ends (epiphyses) of the skeleton suggested that he was an early adolescent with a relatively large body mass, though growth had not yet been completed. On the other hand, study of the dentition, including measurement of microscopic layers of tooth enamel called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1573\">perikymata<\/a><\/strong>, revealed a much younger age of 8 or 9. According to Christopher Dean and Holly Smith (2009), the best explanation for this discrepancy between the dental and skeletal age is that <em>Homo erectus<\/em> had its own distinct growth pattern\u2014reaching maturity more slowly than chimpanzees but faster than <em>Homo sapiens<\/em>. This suggests that the human life history pattern of slow maturation and lengthy dependency was a more recent development. More work remains on refining this pattern for early <em>Homo<\/em>, but it is an important topic that sheds light on how and when we developed our unique life history characteristics.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000;\">The Big Picture of Early <em>Homo<\/em><\/span><\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">We are discovering that the evolution of the genus <em>Homo<\/em> is more complex than what was previously thought. The earlier view of a simple progression from <em>Australopithecus<\/em> to <em>Homo habilis<\/em> to <em>Homo erectus<\/em> as clearly delineated stages in human evolution just doesn\u2019t hold up anymore.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">As is apparent from the information presented here, there is tremendous variability during this time. While fossils classified as <em>Homo habilis<\/em> show many of the characteristics of the genus <em>Homo<\/em>, such as brain expansion and smaller tooth size, the small body size and long arms are more akin to australopithecines. There is also tremendous variability within the fossils assigned to <em>Homo habilis<\/em>, so there is little consensus on whether it is one or multiple species of <em>Homo<\/em>, a member of the genus <em>Australopithecus<\/em>, or even a yet-to-be-defined new genus. Similarly, there are considerable differences in skull morphology and body size and form of <em>Homo erectus<\/em>, of which some specimens show more similarity to <em>Homo habilis<\/em> than previously thought.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">What does this diversity mean for how we should view early <em>Homo<\/em>? First, there isn\u2019t an abrupt break between <em>Australopithecus<\/em> and <em>Homo habilis<\/em> or even between <em>Homo habilis<\/em> and <em>Homo erectus<\/em>. Characteristics we define as <em>Homo<\/em> don\u2019t appear as a unified package; they appear in the fossil record at different times. This is known as <strong>mosaic evolution<\/strong>. Indeed, fossil species such as <em>Australopithecus sediba<\/em>, as well as <em>Homo naledi<\/em> and <em>Homo floresiensis<\/em> (who will be introduced in Chapter 11), have displayed unexpected combinations of primitive and derived traits.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">We can consider several explanations for the diversity we see within early <em>Homo<\/em> from about 2.5 million to 1.5 million years ago. One possibility is the existence of multiple contemporaneous species of early <em>Homo <\/em>during this period. In light of the pattern of environmental instability discussed earlier, it shouldn\u2019t be surprising to see fossils from different parts of Africa and Eurasia display tremendous variability. Multiple hominin forms could also evolve in the same region, as they diversified in order to occupy different ecological niches. However, even the presence of multiple species of hominin does not preclude their interacting and interbreeding with one another. As you\u2019ll see in Appendix D, sequencing of ancient hominin genomes has led to deeper understanding of genetic relationships between extinct species such as the Neanderthals and Denisovans.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Diversity of brain and body sizes could also reflect developmental plasticity\u2014short-term adaptations within a lifetime (Anton et al. 2014). These have the advantage of being more flexible than genetic natural selection, which could only occur over many generations. For example, among human populations today, different body sizes are thought to be adaptations to different climate or nutritional environments. Under Pleistocene conditions of intense variability, a more flexible strategy of adaptation would be valuable.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">New discoveries are also questioning old assumptions about the behavior of <em>Homo habilis<\/em> and <em>Homo erectus<\/em>. Just as the fossil evidence doesn\u2019t neatly separate <em>Australopithecus<\/em> and <em>Homo<\/em>, evidence of the lifeways of early <em>Homo <\/em>show similar diversity. For example, one of the traditional dividing lines between <em>Homo <\/em>and <em>Australopithecus<\/em> was thought to be stone tools: <em>Homo<\/em> made them; <em>Australopithecus <\/em>didn\u2019t. However, the recent discovery of stone tools from Kenya dating to 3.3 million years ago challenges this point of view. Similarly, the belief that <em>Homo erectus<\/em> was the first species to settle outside Africa may now come into question with the report of 2.1-million-year-old stone tools from China. If this find is supported by additional evidence, it may cause a reevaluation of <em>Homo erectus<\/em> being the first to leave Africa. Instead, there could have been multiple earlier migrations of hominins such as <em>Homo habilis<\/em> or even <em>Australopithecus <\/em>species.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">These various lines of evidence about the genus <em>Homo <\/em>point out the need for a more nuanced view of this period of human evolution. Rather than obvious demarcations between species and their corresponding behavioral advancements, it now looks like many behaviors were shared among species. Earlier hominins that we previously didn\u2019t think had the capability could have been doing things like expanding out of Africa or using stone tools. Meanwhile, some other hominins that we had considered more advanced didn\u2019t actually have the full suite of \u201chuman\u201d characteristics previously expected.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">From a student\u2019s perspective, all this complexity probably seems frustrating. It would be ideal if the human story were a straightforward, sequential narrative. Unfortunately, it seems that human evolution was not a nice, neat trajectory of increasingly humanlike traits and behaviors; rather, it is emblematic of the untidy but exciting nature of the study of human evolution.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Despite some haziness dominating the early <em>Homo<\/em> narrative, we can identify some overall trends for the million-year period associated with early <em>Homo. <\/em>These trends include brain expansion, a reduction in facial prognathism, smaller jaw and tooth size, larger body size, and evidence of full terrestrial bipedalism. These traits are associated with a key behavioral shift that emphasizes culture as a flexible strategy to adapt to unpredictable environmental circumstances. Included in this repertoire are the creation and use of stone tools to process meat obtained by scavenging and later hunting, a utilization of fire and cooking, and the roots of the human life history pattern of prolonged childhood, cooperation in child raising, and the practice of skilled foraging techniques. In fact, it\u2019s apparent that the cultural innovations are driving the biological changes, and vice versa, fueling a feedback loop that continues during the later stages of human evolution.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000;\">Hominin Species Summaries<\/span><\/h2>\n<div style=\"text-align: left;\">\n<table class=\"aligncenter\" style=\"width: 344.15pt;\">\n<tbody>\n<tr class=\"Table3-R\" style=\"height: 23pt;\">\n<td class=\"Table3-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><em>Homo habilis <\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 23pt;\">\n<td class=\"Table3-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">2.5 million years ago to 1.7 million years ago<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 23pt;\">\n<td class=\"Table3-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">East and South Africa<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 36pt;\">\n<td class=\"Table3-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Olduvai Gorge, Tanzania; Koobi Fora, Kenya; Sterkfontein, South Africa<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 23pt;\">\n<td class=\"Table3-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">650 cc average (range from 510 cc to 775 cc)<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 23pt;\">\n<td class=\"Table3-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Smaller teeth with thinner enamel compared to <em>Australopithecus<\/em>; parabolic dental arcade shape<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 36pt;\">\n<td class=\"Table3-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Rounder cranium and less facial prognathism than <em>Australopithecus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 36pt;\">\n<td class=\"Table3-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Small stature; similar body plan to <em>Australopithecus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 23pt;\">\n<td class=\"Table3-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Oldowan tools<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 23pt;\">\n<td class=\"Table3-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div style=\"text-align: left;\">\n<table class=\"aligncenter\" style=\"width: 344.15pt;\">\n<tbody>\n<tr class=\"Table4-R\" style=\"height: 23pt;\">\n<td class=\"Table4-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Hominin<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><em>Homo <\/em><em>erectus<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 23pt;\">\n<td class=\"Table4-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Dates<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">1.8 million years ago to about 110,000 years ago<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 23pt;\">\n<td class=\"Table4-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Region(s)<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">East and South Africa; West Eurasia; China and Southeast Asia<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 36pt;\">\n<td class=\"Table4-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Famous discoveries<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Lake Turkana, Olorgesailie, Kenya; Java, Indonesia; Zhoukoudian, China; Dmanisi, Republic of Georgia<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 23pt;\">\n<td class=\"Table4-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Brain size<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Average 900 cc; range between 650 cc and 1,100 cc<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 23pt;\">\n<td class=\"Table4-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Dentition<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Smaller teeth than <em>Homo habilis<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 36pt;\">\n<td class=\"Table4-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Cranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Long, low skull with robust features including thick cranial vault bones and large brow ridge, sagittal keel, and occipital torus<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 36pt;\">\n<td class=\"Table4-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Postcranial features<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Larger body size compared to <em>Homo habilis<\/em>; body proportions (longer legs and shorter arms) similar to <em>Homo sapiens<\/em><\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 23pt;\">\n<td class=\"Table4-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Culture<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Acheulean tools (in Africa); evidence of increased hunting and meat-eating; use of fire; migration out of Africa<\/span><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 23pt;\">\n<td class=\"Table4-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Other<\/strong><\/span><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"padding: 5pt 5pt 5pt 5pt; border: solid #000000 1pt;\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\">N\/A<\/span><\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\"><span style=\"color: #000000;\">Review Questions<strong><br \/>\n<\/strong><\/span><\/h2>\n<ul>\n<li><span style=\"color: #000000;\">Describe the climate during the early Pleistocene. Explain why climate is important for understanding the evolution of early <em>Homo<\/em>.<\/span><\/li>\n<li><span style=\"color: #000000;\">List the key anatomical characteristics that are generally agreed to define the genus <em>Homo<\/em>.<\/span><\/li>\n<li><span style=\"color: #000000;\">Why has classification of early<em> Homo <\/em>fossils proved difficult? What are some explanations for the variability seen in these fossils?<\/span><\/li>\n<li><span style=\"color: #000000;\">Compare and contrast the Oldowan and Acheulean tool industries<em>.<\/em><\/span><\/li>\n<li><span style=\"color: #000000;\">Name some specific behaviors associated with <em>Homo erectus<\/em> in the areas of tool use, subsistence practices, migration patterns, and other cultural innovations.<\/span><\/li>\n<\/ul>\n<\/div>\n<\/div>\n<h2 class=\"__UNKNOWN__\"><span style=\"color: #000000;\">Key Terms<\/span><\/h2>\n<div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Acheulean<\/strong>: Tool industry characterized by teardrop-shaped stone handaxes flaked on both sides. <\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Developmental plasticity<\/strong>: The capability of an organism to modify its phenotype during development in response to environmental cues.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Human behavioral ecology<\/strong>: The study of human behavior from an evolutionary and ecological perspective.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Life history<\/strong>: The broad pattern of a species\u2019 life cycle, including development, reproduction, and longevity.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Mosaic evolution<\/strong>: Different characteristics evolve at different rates and appear at different stages. <\/span><br style=\"clear: both;\" \/><br style=\"clear: both;\" \/><span style=\"color: #000000;\"><strong>Occipital torus<\/strong>: A ridge on the occipital bone in the back of the skull.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Oldowan<\/strong>: Earliest stone-tool industry consisting of simple flakes and choppers.<\/span><br style=\"clear: both;\" \/><span style=\"color: #000000;\"><strong><br style=\"clear: both;\" \/>Perikymata<\/strong>: Microscopic ridges on the surface of tooth enamel that serve as markers of tooth development.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Pleistocene<\/strong>: Geological epoch dating from 2.6 million years ago to about 11,000 years ago.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Pliocene:<\/strong> Geological epoch dating from 5.3 to 2.6 million years ago.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Prognathism<\/strong>: Condition where the lower face and jaw protrude forward from a vertical plane.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><strong>Sagittal keel<\/strong>: A thickened area along the top of the skull.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000;\">About the Author<strong><br \/>\n<\/strong><\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000;\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image14-3.jpg\" alt=\"A woman with long black hair smiles at the camera.\" width=\"222\" height=\"333\" \/><\/span><\/p>\n<h2 class=\"import-Normal\"><strong><span style=\"color: #000000;\">Bonnie Yoshida-Levine, Ph.D.<\/span><\/strong><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Grossmont College, <a class=\"rId108\" style=\"color: #000000;\" href=\"mailto:bonnie.yoshida@gcccd.edu\">bonnie.yoshida@gcccd.edu<\/a><\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Bonnie Yoshida-Levine is an instructor of anthropology at Grossmont College, where she teaches biological anthropology and archaeology. She received her bachelor\u2019s degree in history from the University of California, Los Angeles, and her M.A. and Ph.D. degrees in anthropology from the University of California, Santa Barbara. Her dissertation research focused on the bioarchaeology of early civilizations in north coastal Peru. Bonnie has also collaborated on archaeological field projects in Bolivia and coastal California.<\/span><\/p>\n<\/div>\n<p><span style=\"color: #000000; font-family: Raleway, sans-serif; font-size: 1.5em;\">FOR FURTHER EXPLORATION<\/span><\/p>\n<div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Boaz, Noel Thomas, and Russell L. Ciochon. 2004. <em>Dragon Bone Hill: An Ice-Age Saga of <\/em>Homo erectus. New York: Oxford University Press.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\"><a href=\"https:\/\/humanorigins.si.edu\/\">Human Evolution by the Smithsonian Institution<\/a>.\u00a0Produced by the Smithsonian National Museum of Natural History, this website covers many aspects of human evolution including 3-D models of hominin fossils.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Lewin, Roger, and Robert A. Foley. 2004. <em>Principles of Human Evolution<\/em>. Oxford, UK: Blackwell Publishing.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Mutu, Kari. \u201cHonour Finds Kenya\u2019s Oldest Fossil Hunter Kamoya Kimeu.\u201d <em>The East African<\/em>, July 19, 2021.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Nordling, Linda. \u201cRaising Up African Paleoanthropologists.\u201d <em>SAPIENS, <\/em>September 28, 2021. Accessed February 24, 2023. <a class=\"rId110\" style=\"color: #000000;\" href=\"https:\/\/www.sapiens.org\/biology\/african-paleoanthropologists\/\"><em>https:\/\/www.sapiens.org\/biology\/african-paleoanthropologists\/<\/em><\/a>.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">Risen, Clay. \u201cKamoya Kimeu, Fossil-Hunting \u2018Legend\u2019 in East Africa Is Dead.\u201d<em> New York Times<\/em>, August 11, 2022. Accessed February 24, 2023. https:\/\/www.nytimes.com\/2022\/08\/11\/science\/kamoya-kimeu-dead.html\/.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Stoneking, Mark. 2015. \u201cOf Lice and Men: The Molecular Evolution of Human Lice.\u201d Lecture, Center for Academic Research &amp; Training in Anthropogeny, San Diego, California, October 16, 2015. Accessed February 24, 2023. <a class=\"rId111\" style=\"color: #000000;\" href=\"https:\/\/carta.anthropogeny.org\/events\/unique-features-human-skin\">https:\/\/carta.anthropogeny.org\/events\/unique-features-human-skin<\/a>.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Tarlach, Gemma. 2015. \u201cThe First Humans to Know Winter.\u201d <em>Discover<\/em>, February 26. https:\/\/www.discovermagazine.com\/planet-earth\/the-first-humans-to-know-winter<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Ungar, Peter S. 2017. <em>Evolution's Bite: A Story of Teeth, Diet, and Human Origins<\/em>. Princeton, NJ: Princeton University Press.<strong><br style=\"clear: both;\" \/><\/strong><\/span><\/p>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">References<\/span><\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Aiello, Leslie C., and Peter Wheeler. 1995. \u201cThe Expensive-Tissue Hypothesis.\u201d <em>Current Anthropology<\/em> 36 (2): 199\u2013221.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Anton, Susan C., Richard Potts, and Leslie C. Aiello. 2014. \u201cEvolution of Early <em>Homo<\/em>: An Integrated Biological Perspective.\u201d <em>Science<\/em> 345 (6192) doi: 10.1126\/science.1236828.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Anton, Susan C., and J. Josh Snodgrass. 2012. \u201cOrigins and Evolution of Genus <em>Homo<\/em>: New Perspectives.\u201d <em>Current Anthropology<\/em> 53 (S6): S479\u2013S496.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Ashton, Nick, Simon G. Lewis, Isabelle De Groote, Sarah M. Duffy, Martin Bates, Richard Bates, Peter Hoare, et al. 2014. \u201cHominin Footprints from Early Pleistocene Deposits at Happisburgh, UK.\u201d <em>PLOS ONE<\/em> 9 (2): e88329.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Belmaker, Miriam. 2010. \u201cEarly Pleistocene Faunal Connections between Africa and Eurasia: An Ecological Perspective.\u201d In <em>Out of Africa I: The First Hominin Colonization of Eurasia<\/em>, edited by John G. Fleagle, John J. Shea, Frederick E. Grine, Andrea L. Baden, and Richard E. Leakey, 183\u2013205. Dordrecht: Springer Netherlands.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Blumenschine, Robert, Henry T. Bunn, Valerius Geist, Fumiko Ikawa-Smith, Curtis W. Marean, Anthony G. Payne, John Tooby, J. Nikolaas, and Van Der Merwe. 1987. \u201cCharacteristics of an Early Hominid Scavenging Niche [and Comments and Reply].\u201d <em>Current Anthropology<\/em> 28 (4): 383\u2013407.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Bower, Bruce. 2004. \u201cEvolution\u2019s Buggy Ride.\u201d <em>Science News<\/em> 166 (15): 230\u2013230.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Bramble, Dennis M., and Daniel E. Lieberman. 2004. \u201cEndurance Running and the Evolution of <em>Homo<\/em>.\u201d <em>Nature<\/em> 432 (7015): 345\u2013352.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Coolidge, Frederick L., and Thomas Grant Wynn. 2017. <em>The Rise of Homo Sapiens: The Evolution of Modern Thinking<\/em>. New York: Oxford University Press.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Dean, M. Christopher, and B. Holly Smith. 2009. \u201cGrowth and Development of the Nariokotome Youth, KNM-WT 15000.\u201d In <em>The First Humans\u2013Origin and Early Evolution of the Genus Homo: Contributions from the Third Stony Brook Human Evolution Symposium and Workshop October 3<\/em>\u2013<em>7, 2006<\/em>, edited by Frederick E. Grine, John G. Fleagle, and Richard E. Leakey, 101\u2013120. Dordrecht: Springer Netherlands.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">deMenocal, Peter B. 2014. \u201cClimate Shocks.\u201d <em>Scientific American<\/em> 311 (3): 48\u201353.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Dennell, Robin, and Wil Roebroeks. 2005. \u201cAn Asian Perspective on Early Human Dispersal from Africa.\u201d <em>Nature<\/em> 438 (7071): 1099\u20131104.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Hawkes, Kristen, James F. O\u2019Connell, Nicholas G. Blurton Jones, Helen Alvarez, and Eric L. Charnov. 1998. \u201cGrandmothering, Menopause, and the Evolution of Human Life\u2009Histories.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 95 (3): 1336\u20131339.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Herries, A. I. R., J. M. Martin, A. B. Leece, J. W. Adams, G. Boschian, R. Joannes-Boyau, T. R. Edwards, et al. 2020. \"Contemporaneity of <em>Australopithecus<\/em>, <em>Paranthropus<\/em>, and early <em>Homo erectus<\/em> in South Africa.\" <em>Science<\/em> 368 (6486). https:\/\/doi.org\/10.1126\/science.aaw7293<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Hill, Kim, and Hillard Kaplan. 1999. \u201cLife History Traits in Humans: Theory and Empirical Studies.\u201d <em>Annual Review of Anthropology<\/em> 28: 397\u2013430.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Hlubik, Sarah, Francesco Berna, Craig Feibel, David Braun, and John W. K. Harris. 2017. \u201cResearching the Nature of Fire at 1.5 Mya on the Site of FxJj20 AB, Koobi Fora, Kenya, Using High-Resolution Spatial Analysis and FTIR Spectrometry.\u201d <em>Current Anthropology<\/em> 58 (S16): S243\u2013S257.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Jablonski, Nina G. 2010. \u201cThe Naked Truth.\u201d <em>Scientific American<\/em> 302 (2): 42\u201349.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Kamberov, Yana G., Elinor K. Karlsson, Gerda L. Kamberova, Daniel E. Lieberman, Pardis C. Sabeti, Bruce A. Morgan, and Clifford J. Tabin. 2015. \u201cA Genetic Basis of Variation in Eccrine Sweat Gland and Hair Follicle Density.\u201d <em>Proceedings of the National Academy of Sciences<\/em> 112 (32): 9932\u20139937.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff; color: #ffffff; margin-left: 0pt; margin-right: 4pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Kittler, R., M. Kayser, and M. Stoneking. 2003. \"Molecular Evolution of Pediculus Humanus and the Origin of Clothing.\" <em>Current Biology<\/em> 13 (16): 1414\u20131417.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Leakey, Louis S. B., Phillip V. Tobias, and John R. Napier. 1964. \u201cA New Species of Genus <em>Homo<\/em> from Olduvai Gorge.\u201d <em>Nature<\/em> 202: 308\u2013312.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Lemorini, Cristina, Thomas W. Plummer, David R. Braun, Alyssa N. Crittenden, Peter W. Ditchfield, Laura C. Bishop, Fritz Hertel, et al. 2014. \u201cOld Stones\u2019 Song: Use-Wear Experiments and Analysis of the Oldowan Quartz and Quartzite Assemblage from Kanjera South (Kenya).\u201d <em>Journal of Human Evolution<\/em> 72: 10\u201325.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Lisiecki, Lorraine E., and Maureen E. Raymo. 2005. \"A Pliocene-Pleistocene stack of 57 globally distributed benthic \u03b418O records.\" <em>Paleoceanography<\/em> 20 (1)<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Lordkipanidze, David, Marcia S. Ponce de Le\u00f3n, Ann Margvelashvili, Yoel Rak, G. Philip Rightmire, Abesalom Vekua, and Christoph P. E. Zollikofer. 2013. \u201cA Complete Skull from Dmanisi, Georgia, and the Evolutionary Biology of Early <em>Homo<\/em>.\u201d <em>Science<\/em> 342 (6156): 326\u2013333.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Matsu'ura, S., M. Kondo, T. Danhara, S. Sakata, H. Iwano, T. Hirata, I. Kurniawan, et al. 2020. \"Age Control of the First Appearance Datum for Javanese <em>Homo erectus<\/em> in the Sangiran Area.\" <em>Science<\/em> 367 (6474): 210\u2013214.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Qiu, Jane. 2016. \u201cHow China Is Rewriting the Book on Human Origins.\u201d <em>Nature<\/em> 535: 22\u201325.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff; color: #ffffff; margin-left: 0pt; margin-right: 4pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Reed, David L., Jessica E. Light, Julie M. Allen, and Jeremy J. Kirchman. 2007. \"Pair of Lice Lost or Parasites Regained: The Evolutionary History of Anthropoid Primate Lice.\" <em>BMC Biology<\/em> 5 (1): 7. doi: 10.1186\/1741-7007-5-7.<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff; color: #ffffff; margin-left: 0pt; margin-right: 4pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Rizal, Y., K. E. Westaway, Y. Zaim, G. D. van den Bergh, E. A. Bettis, 3rd, M. J. Morwood, O. F. Huffman, R. Gr\u00fcn, et al. 2020. \"Last Appearance of <em>Homo erectus<\/em> at Ngandong, Java, 117,000\u2013108,000 Years Ago.\" <em>Nature<\/em> 577 (7790): 381\u2013385.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Roche, Helene, Robert J. Blumenschine, and John J. Shea. 2009. \u201cOrigins and Adaptations of Early <em>Homo<\/em>: What Archeology Tells Us.\u201d In <em>The First Humans: Origin and Early Evolution of the Genus Homo<\/em>, edited by Frederick E. Grine, John G. Fleagle, and Richard E. Leakey, 135\u2013147. New York: Springer.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Rogers, Alan R., David Iltis, and Stephen Wooding. 2004. \u201cGenetic Variation at the MC1R l Locus and the Time since Loss of Human Body Hair.\u201d <em>Current Anthropology<\/em> 45 (1): 105\u2013108.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Ruff, Christopher. 2009. \u201cRelative Limb Strength and Locomotion in <em>Homo<\/em><em>habilis<\/em>.\u201d <em>American Journal of Physical Anthropology<\/em> 138 (1): 90\u2013100.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Shipton, Ceri, James Blinkhorn, Paul S. Breeze, Patrick Cuthbertson, Nick Drake, Huw S. Groucutt, Richard P. Jennings, et al. 2018. \u201cAcheulean Technology and Landscape Use at Dawadmi, Central Arabia.\u201d <em>PloS one<\/em> 13 (7): e0200497\u2013e0200497.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Simpson, Scott W., Jay Quade, Naomi E. Levin, Robert Butler, Guillaume Dupont-Nivet, Melanie Everett, and Sileshi Semaw. 2008. \u201cA Female <em>Homo<\/em><em>erectus<\/em> Pelvis from Gona, Ethiopia.\u201d <em>Science<\/em> 322 (5904): 1089\u20131092.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Ungar, Peter S., and Robert S. Scott. 2009. \u201cDental Evidence for Diets of Early <em>Homo<\/em>.\u201d In <em>The First Humans: Origin and Early Evolution of the Genus Homo<\/em>, edited by Frederick E. Grine, John G. Fleagle, and Richard E. Leakey, 121\u2013134. New York: Springer.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Villmoare, Brian, William H. Kimbel, Chalachew Seyoum, Christopher J. Campisano, Erin N. DiMaggio, John Rowan, David R. Braun, J. Ram\u00f3n Arrowsmith, and Kaye E. Reed. 2015. \u201cEarly <em>Homo<\/em> at 2.8 Ma From Ledi-Geraru, Afar, Ethiopia.\u201d <em>Science<\/em> 347 (6228): 1352\u20131355.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Wood, Bernard. 2014. \u201cHuman Evolution: Fifty Years after <em>Homo<\/em><em>habilis<\/em>.\u201d <em>Nature<\/em> 508 (7494): 31\u201333.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Wood, Bernard, and Mark Collard. 1999. \u201cThe Changing Face of Genus <em>Homo<\/em>.\u201d <em>Evolutionary Anthropology<\/em> 8 (6): 195\u2013207.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Wrangham, Richard. 2009. <em>Catching Fire: How Cooking Made Us Human<\/em>. New York: Basic Books.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt; margin-right: -36pt; text-indent: 0pt;\"><span style=\"color: #000000;\">Zhu, Zhaoyu, Robin Dennell, Weiwen Huang, Yi Wu, Shifan Qiu, Shixia Yang, and Zhiguo Rao. 2018. \u201cHominin Occupation of the Chinese Loess Plateau Since about 2.1 Million Years Ago.\u201d <em>Nature<\/em> 559: 608\u2013612.<\/span><\/p>\n<h2 class=\"import-Normal\"><span style=\"color: #000000;\">Acknowledgments<br \/>\n<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"color: #000000;\">The author gratefully acknowledges funding from the California Community Colleges Chancellor\u2019s Office Zero Textbook Cost Degree Grant Program\u2014Implementation Phase 2.<\/span><\/p>\n<\/div>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1775\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1775\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\">Jonathan Marks, Ph.D., University of North Carolina at Charlotte<\/p>\n<p class=\"import-Normal\">Adam P. Johnson, M.A., University of North Carolina at Charlotte\/University of Texas at San Antonio<\/p>\n<p class=\"import-Normal\"><em>This chapter is an adaptation of \"<\/em><a class=\"rId9\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__\/\"><em>Chapter 2: Evolution<\/em><\/a><em>\u201d by Jonathan Marks. In <\/em><a class=\"rId10\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, first edition<\/em><\/a><em>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under <\/em><a class=\"rId11\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\"><em>CC BY-NC 4.0<\/em><\/a><em>. <\/em><\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\"><span style=\"color: #ffffff\">Learning Objectives<\/span><\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li>Explain the relationship among genes, bodies, and organismal change.<\/li>\n<li>Discuss the shortcomings of simplistic understandings of genetics.<\/li>\n<li>Describe what is meant by the \"biopolitics of heredity.\"<\/li>\n<li>Discuss issues caused by misuse of ideas about adaptations and natural selection.<\/li>\n<li>Examine and correct myths about evolution.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p class=\"import-Normal\">The Human Genome Project, an international initiative launched in 1990, sought to identify the entire genetic makeup of our species. For many scientists, it meant trying to understand the genetic underpinnings of what made humans uniquely human. James Watson, a codiscoverer of the helical shape of DNA, wrote that \u201cwhen finally interpreted, the genetic messages encoded within our DNA molecules will provide the ultimate answers to the chemical underpinnings of human existence\u201d (Watson 1990, 248). The underlying message is that what makes humans unique can be found in our <strong>genes<\/strong>. The Human Genome Project hoped to find the core of who we are and where we come from.<\/p>\n<p class=\"import-Normal\">Despite its lofty goal, the Human Genome Project\u2014even after publishing the entire human genome in January 2022\u2014could not fully account for the many factors that contribute to what it is to be human. Richard Lewontin, Steven Rose, and Leon Kamin (2017) argue that genetic determinism of the sort assumed by the Human Genome Project neglects other essential dimensions that contribute to the development and evolution of human bodies, not to mention the role that culture plays. They use an apt metaphor of a cake to illustrate the incompleteness of reductive models. Consider the flavor of a cake and think of the ingredients listed in the recipe. The recipe includes ingredients such as flour, sugar, shortening, vanilla extract, eggs, and milk. Does raw flour taste like cake? Does sugar, vanilla extract, or any of the other ingredients taste like cake? They do not, and knowing the individual flavors of each ingredient does not tell us much about what cake tastes like. Even mixing all of the ingredients in the correct proportions does not get us cake. Instead, external factors such as baking at the right temperature, for the right amount of time, and even the particularities of our evolved sense of taste and smell are all necessary components of experiencing the cake.<\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ff00ff\">Lewontin, Rose, and Kamin (2017) argue that the same is true for humans and other organisms.<\/span><\/p>\n<p class=\"import-Normal\">Knowing everything about cake ingredients does not allow us to fully know cake. Equally so, knowing everything about the genes found in our DNA does not allow us to fully know humans. Different, interacting levels are implicated in the development and evolution of all organisms, including humans. Genes, the structure of chromosomes, developmental processes, epigenetic tags, environmental factors, and still-other components all play key roles such that genetically reductive models of human development and evolution are woefully inadequate.<\/p>\n<p class=\"import-Normal\">The complex interactions across many levels\u2014genetic, developmental, and environmental\u2014explain why we still do not know how our one-dimensional DNA nucleotide sequence results in a four-dimensional organism. This was the unfulfilled promise of the inception of the Human Genome Project in the 1980s and 1990s: the project produced the complete DNA sequence of a human cell in the hopes that it would reveal how human bodies are built and how to cure them when they are built poorly. Yet, that information has remained elusive. Presumably, the knowledge of how organisms are produced from DNA sequences will one day permit us to reconcile the discrepancies between patterns in anatomical evolution and molecular evolution.<\/p>\n<p class=\"import-Normal\">In this chapter, we will consider multilevel evolution and explore evolution as a complex interaction between genetic and epigenetic factors as well as the environments in which organisms live. Next, we will examine the biopolitical nature of human evolution. We will then investigate problems that arise from attributing all traits to an adaptive function. Finally, we will address common misconceptions about evolution. The goal of this chapter is to provide you with the necessary toolkit for understanding the molecular, anatomical, and political dimensions of evolution.<\/p>\n<h2 class=\"import-Normal\">Evolution Happens at Multiple Levels<\/h2>\n<p class=\"import-Normal\">Following Richard Dawkins\u2019s publication of <em>The Selfish Gene <\/em>in 1976, the scientific imagination was captured by the potential of genomics to reveal how genes are copied by Darwinian selection. Dawkins argues that the genes in individuals that contribute to greater reproductive success are the units of selection. His conception of evolution at the molecular level undercuts the complex interactions between organisms and their environments, which are not expressed genomically but are nevertheless key drivers in evolution.<\/p>\n<p class=\"import-Normal\">By the 1980s, the acknowledgment among most biologists that even though genes construct bodies, genes and bodies evolve at different rates and with distinct patterns. This realization led to a renewed focus on how bodies change. The Evolutionary Synthesis of the 1930s\u20131970s had reduced organisms to their <strong>genotypes<\/strong> and species to their <strong>gene pools<\/strong>, which provided valuable insights about the processes of biological change, but it was only a first approximation. Animals are in fact reactive and adaptable beings, not passive and inert genotypes. Species are clusters of socially interacting and reproductively compatible organisms.<\/p>\n<figure style=\"width: 291px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/06\/image8-5.png\" alt=\"An asteroid hits the ocean. Pterodactyls fly among clouds in the foreground.\" width=\"291\" height=\"233\" \/><figcaption class=\"wp-caption-text\">Figure 17.1: A painting by Donald E. Davis representing the Chicxulub asteroid impact off the Yucatan Peninsula that contributed to the mass extinction that included the dinosaurs about 65 million years ago. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Chicxulub_impact_-_artist_impression.jpg\">Chicxulub impact - artist impression<\/a> by Donald E. Davis, <a href=\"https:\/\/www.nasa.gov\/\">NASA<\/a>, is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Once we accept that evolutionary change is fundamentally genetic change, we can ask: How do bodies function and evolve? How do groups of animals come to see one another as potential mates or competitors for mates, as opposed to just other creatures in the environment? Are there evolutionary processes that are not explicable by population genetics? These questions\u2014which lead us beyond reductive assumptions\u2014were raised in the 1980s by Stephen Jay Gould, the leading evolutionary biologist of the late 20th century (see: Gould 2003; 1996).<\/p>\n<p class=\"import-Normal\">Gould spearheaded a movement to identify and examine higher-order processes and features of evolution that were not adequately explained by population genetics. For example, <strong>extinction<\/strong>, which was such a problem for biologists of the 1600s, could now be seen as playing a more complex role in the history of life than population genetics had been able to model. Gould recognized that there are two kinds of extinctions, each with different consequences: background extinctions and mass extinctions. Background extinctions are those that reflect the balance of nature, because in a competitive Darwinian world, some things go extinct and other things take their place. Ecologically, your species may be adapted to its niche, but if another species comes along that\u2019s better adapted to the same niche, eventually your species will go extinct. It sucks, but it is the way of all life: you come into existence, you endure, and you pass out of existence. But mass extinctions are quite different. They reflect not so much the balance of nature as the wholesale disruption of nature: many species from many different lineages dying off at roughly the same time\u2014presumably as the result of some kind of rare ecological disaster. The situation may not be survival of the fittest as much as survival of the luckiest. The result, then, would be an ecological scramble among the survivors. Having made it through the worst, the survivors could now simply divide up the new ecosystem amongst themselves, since their competitors were gone. Something like this may well have happened about 65 million years ago, when a huge asteroid hit the Yucatan Peninsula, which mammals survived but dinosaurs did not (Figure 17.1). Something like this may be happening now, due to human expansion and environmental degradation. Note, though, that there is only a limited descriptive role here for population genetics: the phenomena we are describing are about organisms and species in ecosystems.<\/p>\n<p class=\"import-Normal\">Another question involved the disconnect between properties of <em>species<\/em> and the properties of <em>gene pools<\/em>. For example, there are upwards of 15 species of gibbons but only two species of chimpanzees. Why? There are upwards of 20 species of guenons but fewer than ten of baboons. Why? Are there genes for that? It seems unlikely. Gould suggested that species, as units of nature, might have properties that are not reducible to the genes in their cells. For example, rates of speciation and extinction might be properties of their ecologies and histories rather than their genes. Thus, relationships between environmental contexts and variability within a species result in degrees of resistance to extinction and affect the frequency and rates at which clades diversify (Lloyd and Gould 1993). Consistent biases of speciation rates might well produce patterns of macroevolutionary diversity that are difficult to explain genetically and better understood ecologically. Gould called such biases in speciation rates <strong>species selection<\/strong>\u2014a higher-order process that invokes competition between species, in addition to the classic Darwinian competition between individuals.<\/p>\n<p class=\"import-Normal\">One of Gould\u2019s most important studies involved the very nature of species. In the classical view, a species is continually adapting to its environment until it changes so much that it is a different species than it was at the beginning of this sentence (Eldredge and Gould 1972). That implies that the species is a fundamentally unstable entity through time, continuously changing to fit in. But suppose, argued Gould along with paleontologist Niles Eldredge, a species is more stable through time and only really adapts during periods of ecological instability and change. Then we might expect to find in the fossil record long equilibrium periods\u2014a few million years or so\u2014in which species don\u2019t seem to change much, punctuated by relatively brief periods in which they change a bit and then stabilize again as new species. They called this idea <strong>punctuated equilibria<\/strong>. The idea helps to explain certain features of the fossil record, notably the existence of small anatomical \u201cgaps\u201d between closely related fossil forms (Figure 17.2). Its significance lies in the fact that although it incorporates genetics, punctuated equilibria is not really a theory of genetics but one of types bodies in deep time.<\/p>\n<p class=\"import-Normal\">Punctuated equilibria is seen across taxa, with long periods in the fossil record representing little phenotypic change. These periods of stability are disrupted by shorter periods of rapid <strong>adaptation<\/strong>, the process through which populations of organisms become suited to living in their environments. Phenotypic changes are often coupled with drastic climatic or ecological changes that affect the milieu in which organisms live. For example, throughout much of hominin evolutionary history, brain size was closely associated with body size and thus remained mostly stable. However, changes occurred in average hominin brain size at around 100 thousand years ago, 1 million years ago, and 1.8 million years ago. Several hypotheses have been put forth to explain these changes, including unpredictability in climate and environment (Potts 1998), social development (Barton 1996), and the evolution of language (Deacon 1998). Evidence from the fossil record, paleoclimate models, and comparative anatomy suggests that the changes observed in hominin lineage result from biocultural processes\u2014that is, the coalescence of environmental and cultural factors that selected for larger brains (Marks 2015; Shultz, Nelson, and Dunbar 2012).<\/p>\n<figure style=\"width: 461px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image5-8.png\" alt=\"Two graphs contrast phyletic gradualism and punctuated equilibria.\" width=\"461\" height=\"222\" \/><figcaption class=\"wp-caption-text\">Figure 17.2: Different ways of conceptualizing the evolutionary relationship between an earlier and a later species. With phyletic gradualism, species are envisioned transforming continually in a direct line over time. With punctuated equilibria species branch off at particular points over time.\u00a0 Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__\/\">Phyletic gradualism vs. punctuated equilibria (Figure 2.12)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">In response to the call for a theory of the evolution of form, the field of <strong>evo-devo<\/strong>\u2014the intersection of evolutionary and developmental biology\u2014arose. The central focus here is on how changes in form and shape arise. An embryo matures by the stimulation of certain cells to divide, forming growth fields. The interactions and relationships among these growth fields generate the structures of the body. The <strong>hox genes<\/strong> that regulate these growth fields turn out to be highly conserved across the animal kingdom. This is because they repeatedly turn on and off the most basic genes guiding the animal\u2019s development, and thus any changes to them would be catastrophic. Indeed, these genes were first identified by manipulating them in fruit flies, such that one could produce a bizarre mutant fruit fly that grew a pair of legs where its antennae were supposed to be (Kaufman, Seeger, and Olsen 1990).<\/p>\n<p class=\"import-Normal\">Certain genetic changes can alter the fates of cells and the body parts, while other genetic changes can simply affect the rates at which neighboring groups of cells grow and divide, thus producing physical bumps or dents in the developing body. The result of altering the relationships among these fields of cellular proliferation in the growing embryo is <strong>allometry<\/strong>, or the differential growth of body parts. As an animal gets larger\u2014either over the course of its life or over the course of macroevolution\u2014it often has to change shape in order to live at a different size. Many important physiological functions depend on properties of geometric area: the strength of a bone, for example, is proportional to its cross-sectional area. But area is a two-dimensional quality, while growing takes place in three dimensions\u2014as an increase in mass or volume. As an animal expands, its bones necessarily weaken, because volume expands faster than area does. Consequently a bigger animal has more stress on its bones than a smaller animal does and must evolve bones even thicker than they would be by simply scaling the animal up proportionally. In other words, if you expand a mouse to the size of an elephant, it will nevertheless still have much thinner bones than the elephant does. But those giant mouse bones will unfortunately not be adequate to the task. Thus, a giant mouse would have to change aspects of its form to maintain function at a larger size (see Figure 17.3).<\/p>\n<p class=\"import-Normal\"><img class=\"aligncenter\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image6-6.png\" alt=\"Side-view of a mouse skeleton.\" width=\"515\" height=\"252\" \/><\/p>\n<figure style=\"width: 453px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image7-9.png\" alt=\"Side-view of an elephant skeleton.\" width=\"453\" height=\"326\" \/><figcaption class=\"wp-caption-text\">Figure 17.3: Mouse (top) and elephant (bottom) skeletons. Notice the elephant\u2019s bones are more robust when the two animals are the same size. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__\/\">Mouse and elephant skeletons (Figure 2.13)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Physiologically, we would like to know how the body \u201cknows\u201d when to turn on and off the genes that regulate growth to produce a normal animal. Evolutionarily, we would like to know how the body \u201clearns\u201d to alter the genetic on\/off switch (or the genetic \u201cslow down\/speed up\u201d switch) to produce an animal that looks different. Moreover, since organisms differ from one another, we would like to know how the developing body distinguishes a range of normal variation from abnormal variation. And, finally, how does abnormal variation eventually become normal in a descendant species?<\/p>\n<p class=\"import-Normal\">Taking up these questions, Gould invoked the work of a British geneticist named Conrad H. Waddington, who thought about genetics in less reductive ways than his colleagues. Rather than isolate specific DNA sites to analyze their function, Waddington instead studied the inheritance of an organism\u2019s reactivity\u2014its ability to adapt to the circumstances of its life. In a famous experiment, he grew fruit fly eggs in an atmosphere containing ether. Most died, but a few survived somehow by developing a weird physical feature: a second thorax with a second pair of wings. Waddington bred these flies and soon developed a stable line of flies who would reliably develop a second thorax when grown in ether. Then he began to lower the concentration of ether, while continuing to selectively breed the flies that developed the strange appearance. Eventually he had a line of flies that would stably develop the \u201cbithorax\u201d <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1238\"><strong>phenotype<\/strong><\/a>\u2013the suite of traits of an organism\u2013even when there was no ether; it had become the \u201cnew normal.\u201d The flies had genetically assimilated the bithorax condition.<\/p>\n<p class=\"import-Normal\">Waddington was thus able to mimic the <strong>inheritance of acquired characteristics<\/strong>: what had been a trait stimulated by ether a few generations ago was now a normal part of the development of the descendants. Waddington recognized that while he had performed a selection experiment on genetic variants, he had not selected for particular traits. Rather, he helped produce the physiological tendency to develop particular traits when appropriately stimulated. He called that tendency <strong>plasticity<\/strong> and its converse, the tendency to stay the same even under weird environmental circumstances, <strong>canalization.<\/strong> Waddington had initially selected for plasticity, the tendency to develop the bithorax phenotype under weird conditions, and then, later, for canalization, the developmental normalization of that weird physical trait. Although Waddington had high stature in the community of geneticists, evolutionary biologists of the 1950s and 1960s regarded him with suspicion because he was not working within the standard mindset of reductionism, which saw evolution as the spread of genetic variants that coded for favorable traits. Both Waddington and Gould resisted contemporary intellectual paradigms that favored reductive accounts of evolutionary processes. They conceived of evolution as an emergent process in which many external factors (e.g. climate, environment, predation) and internal factors (e.g., genotypes, plasticity, canalization) coalesce to produce the evolutionary trends that we observe in the fossil record and our genome.<\/p>\n<p class=\"import-Normal\">While Gould and Waddington both looked beyond the genome to understand evolution, the Human Genome Project\u2014an international project with the goal of identifying each base pair in the human genome in the 1990s\u2014generated a great deal of public interest in analyzing the human DNA sequence from the standpoint of medical genetics. Some of the rhetoric aimed to sell the public on investing a lot of money and resources in sequencing the human genome in order to show the genetic basis of heritable traits, cure genetic diseases, and learn what it means ultimately to be biologically human. However, the Human Genome Project was not actually able to answer those questions through the use of genetics alone, and thus a broader, more holistic account was required.<\/p>\n<p class=\"import-Normal\">This holistic account came from decades of research in human biology and anthropology, which understood the human body as highly adaptable, dynamic, and emergent. For example, in the early 20th century, anthropologist Franz Boas measured the skulls of immigrants to the U.S., revealing that environmental, not merely genetic, factors affected skull shape. The growing human body adjusts itself to the conditions of life, such as diet, sunshine, high altitude, hard labor, population density, how babies are carried\u2014any and all of which can have subtle but consistent effects upon its development. There can thus be no normal human form, only a context-specific range of human forms.<\/p>\n<p class=\"import-Normal\">However, what the human biologists called human adaptability, evolutionary biologists called developmental plasticity, and evidence quickly began to mount for its cause being <strong>epigenetic <\/strong>modifications to DNA. Epigenetic modifications are changes to how genes are used by the body (as opposed to changes in the DNA sequences; see Chapter 3). Scientific interest shifted from the focus of the Human Genome Project to the ways that bodies are made by evolutionary-developmental processes, including epigenetics. What is meant by \u201cepigenetic modification\u201d? Evolution is about how descendants diverge from their ancestors. Inheritance from parent to offspring is still critical to this process, which occurs through genetic recombination: the pairing of homologous chromosomes and sharing of genetic material during meiosis (see Chapter 3). However, in the 21st century, the link between evolution and inheritance has broadened with a clearer understanding of how environmental and developmental factors shape bodies and the expression of genes, including epigenetic inheritance patterns. While offspring inherit their genes through random assortment during meiosis, environmental factors also shape how genes are used. When these epigenetic modifications occur in germ cells, they can be passed onto offspring. In these cases, there is no change in the DNA sequence but rather in how genes are used by the body due to DNA methylation and the structure of chromosomes due to histone acetylation (see Chapter 3).<\/p>\n<p class=\"import-Normal\">In addition, we now recognize that evolution is affected by two other forms of intergenerational transmission and inheritance (in addition to genetics and epigenetics). These forms include behavioral variation and culture. That is, behavioral information can be transmitted horizontally (intragenerationally), permitting more rapid ways for organisms to adjust to the environment. And, then there is the fourth mode of transmission: the cultural or symbolic mode. <span style=\"background-color: #ffff00\">Humans are the only species<\/span> that horizontally transmits an arbitrary set of rules to govern communication, social interaction, and thought. This shared information is symbolic and has resulted in what we recognize as \u201cculture\u201d: locally emergent worlds of names, words, pictures, classifications, revered pasts, possible futures, spirits, dead ancestors, unborn descendants, in-laws, politeness, taboo, justice, beauty, and story, all accompanied by practices and a material world of tools.<\/p>\n<p class=\"import-Normal\">Consequently our contemporary ideas about evolution see the evolutionary processes as hierarchically organized and not restricted to the differential transmission of DNA sequences into the next generation. While that is indeed a significant part of evolution, the organism and species are nevertheless crucial to understanding how those DNA sequences get transmitted. Further, the transmission of epigenetic, behavioral, and symbolic information play a complex role in perpetuating our genes, bodies, and species. In the case of human evolution, one can readily see that symbolic information and cultural adaptation are far more central to our lives and our survival today than DNA and genetic adaptation. It is thus misleading to think of humans passively occupying an environmental niche. Rather, humans are actively engaged in constructing our own niches, as well as adapting to them and using them to adapt. The complex interplay between a species and its active engagement in creating its own ecology is known as <strong>niche construction<\/strong>. If we understand <strong>natural selection<\/strong>\u2013the process by which populations adapt to their specific environments\u2013as the effects that environmental context has on the reproductive success of organisms, then niche construction is the process through which organisms shape their own selective pressures.<\/p>\n<h2 class=\"import-Normal\">The Biopolitics of Heredity<\/h2>\n<p class=\"import-Normal\">\u201cScience isn\u2019t political\u201d is a sentiment that you have likely heard before. Science is supposed to be about facts and objectivity. It exists, or at least ought to, outside of petty human concerns. However, the sorts of questions we ask as scientists, the problems we choose to study, the categories and concepts we use, who gets to do science, and whose work gets cited are all shaped by culture. Doing science is a political act. This fact is markedly true for human evolution. While it is easier to create intellectual distance between us and fruit flies and viruses, there is no distance when we are studying ourselves. The hardest lesson to learn about human evolution is that it is intensely political. Indeed, to see it from the opposite side, as it were, the history of creationism\u2014the belief that the universe was divinely created around 6,000 years ago\u2014is essentially a history of legal decisions. For instance, in <em>Tennessee v. John T. Scopes<\/em> (1925), a schoolteacher was prosecuted for violating a law in Tennessee that prohibited the teaching of human evolution in public schools, where teachers were required by law to teach creationism.<\/p>\n<p class=\"import-Normal\">More recently, legal decisions aimed at legislating science education have shaped how students are exposed to evolutionary theory. For instance, <em>McLean v. Arkansas<\/em> (1982) dispatched \u201cscientific creationism\u201d by arguing that the imposition of balanced teaching of evolution and creationism in science classes violates the Establishment Clause, separating church and state. Additionally, <em>Kitzmiller v. Dover (Pennsylvania) Area School District<\/em> (2005) dispatched the teaching of \u201cintelligent design\u201d in public school classrooms as it was deemed to not be science. In some cases, people see unbiblical things in evolution, although most Christian theologians are easily able to reconcile science to the Bible. In other cases, people see immoral things in evolution, although there is morality and immorality everywhere. And some people see evolution as an aspect of alt-religion, usurping the authority of science in schools to teach the rejection of the Christian faith, which would be unconstitutional due to the protected separation of church and state.<\/p>\n<p class=\"import-Normal\">Clearly, the position that politics has nothing to do with science is untenable. But is the politics in evolution an aberration or is it somehow embedded in science? In the early 20th century, scientists commonly promoted the view that science and politics were separate: science was seen as a pure activity, only rarely corrupted by politics. And yet as early as World War I, the politics of nationalism made a hero of the German chemist Fritz Haber for inventing poison gas. And during World War II, both German doctors and American physicists, recruited to the war effort, helped to end many civilian lives. Therefore, we can think of the apolitical scientist as a self-serving myth that functions to absolve scientists of responsibility for their politics. The history of science shows how every generation of scientists has used evolutionary theory to rationalize political and moral positions. In the very first generation of evolutionary science, Darwin\u2019s <em>Origin of Species<\/em> (1859) is today far more readable than his <em>Descent of Man<\/em> (1871). The reason is that Darwin consciously purged <em>The Origin of Species<\/em> of any discussion of people. And when he finally got around to talking about people, in <em>The Descent of Man<\/em>, he simply imbued them with the quaint Victorian prejudices of his age, and the result makes you cringe every few pages. There is plenty of politics in there\u2014sexism, racism, and colonialism\u2014because <em>you cannot talk about people apolitically<\/em>.<\/p>\n<p class=\"import-Normal\">One immediate faddish deduction from Darwinism, popularized by Herbert Spencer (1864) as \u201csurvival of the fittest,\u201d held that unfettered competition led to advancement in nature and to human history. Since the poor were purported losers in that struggle, anything that made their lives easier would go against the natural order. This position later came to be known ironically as \u201cSocial Darwinism.\u201d Spencer was challenged by fellow Darwinian Thomas Huxley (1863), who agreed that struggle was the law of the jungle but observed that we don\u2019t live in jungles anymore. The obligation to make lives better for others is a moral, not a natural, fact. We simultaneously inhabit a natural universe of descent from apes and a moral universe of injustice and inequality, and science is not well served by ignoring the latter.<\/p>\n<p class=\"import-Normal\">Concurrently, the German biologist Ernst Haeckel\u2019s 1868 popularization of Darwinism was translated into English a few years later as <em>The History of Creation<\/em>. As we saw earlier, Haeckel was determined to convince his readers that they were descended from apes, even in the absence of fossil evidence attesting to it. When he made non-Europeans into the missing links that connected his readers to the apes, and depicted them as ugly caricatures, he knew precisely what he was doing. Indeed, even when the degrading racial drawings were deleted from the English translation of his book, the text nevertheless made his arguments quite clear. And a generation later, when the Americans had not yet entered the Great War in 1916, a biologist named Vernon Kellogg visited the German High Command as a neutral observer and found that the officers knew a lot about evolutionary biology, which they had gotten from Haeckel and which rationalized their military aggressions. Kellogg went home and wrote a bestseller about it, called <em>Headquarters Nights<\/em> (1917). World War I would have been fought with or without evolutionary theory, but as a source of scientific authority, evolution\u2014even if a perversion of the Darwinian theory\u2014had very quickly attained global geopolitical relevance.<\/p>\n<p class=\"import-Normal\">Oftentimes, politics in evolutionary science is subtle, due to the pervasive belief in the advancement of science. We recognize the biases of our academic ancestors and modify our scientific stories accordingly. But we can never be free of our own cultural biases, which are invisible to us, as much as our predecessors\u2019 biases were invisible to them. In some cases, the most important cultural issues resurface in different guises each generation, like scientific racism. <strong>Scientific racism<\/strong> is the recruitment of science for the evil political ends of racism, and it has proved remarkably impervious to evolution. Before Darwin, there was creationist scientific racism, and after Darwin, there was evolutionist scientific racism. And there is still scientific racism today, self-justified by recourse to evolution, which means that scientists have to be politically astute and sensitive to the uses of their work to counter these social tendencies.<\/p>\n<p class=\"import-Normal\">Consider this: Are you just your ancestry, or can you transcend it? If that sounds like a weird question, it was actually quite important to a turn-of-the-20th-century European society in which an old hereditary aristocracy was under increasing threat from a rising middle class. And that is why the very first English textbook of Mendelian genetics concluded with the thought that \u201cpermanent progress is a question of breeding rather than of pedagogics; a matter of gametes, not of training \u2026 the creature is not made but born\u201d (Punnett 1905, 60). <em>Translation: Not only do we now know a bit about how heredity works, but it\u2019s also the most important thing about you. Trust me, I\u2019m a scientist.<\/em><\/p>\n<p class=\"import-Normal\">Yet evolution is about how descendants come to differ from ancestors. Do we really know that your heredity, your DNA, your ancestry, is the most important thing about you? That you were born, not made? After all, we do know that you could be born into slavery or as a peasant, and come from a long line of enslaved people or peasants, and yet not have slavery or peasantry be the most important thing about you. Whatever your ancestors were may unfortunately constrain what you can become, but as a moral precept, it should not. But just as science is not purely \u201cfacts and objectivity,\u201d ancestry is not a strictly biological concept. Human ancestry is biopolitics, not biology.<\/p>\n<p class=\"import-Normal\">Evolution is fundamentally a theory about ancestry, and yet ancestors are, in the broad anthropological sense, sacred: ancestors are often more meaningful symbolically than biologically. Just a few years after <em>The Origin of Species <\/em>(Darwin 1859), the British politician and writer Benjamin Disraeli declared himself to be on the side of the angels, not the apes, and to \u201crepudiate with indignation and abhorrence those new-fangled theories\u201d (Monypenny, Flavelle, and Buckle 1920, 105). He turned his back on an ape ancestry and looked to the angel; yet, he did so as a prominent Jew-turned-Anglican, who had personally transcended his humble roots and risen to the pinnacle of the Empire. Ancestry was certainly important, and Disraeli was famously proud of his, but it was also certainly not the most important thing, not the primary determinant of his place in the world. Indeed, quite the opposite: Disraeli\u2019s life was built on the transcendence of many centuries of Jewish poverty and oppression in Europe. Humble ancestry was there to be superseded and nobility was there to be earned; Disraeli would later become the Earl of Beaconsfield. Clearly, \u201care you just your ancestry\u201d is not a value-neutral question, and \u201cthe creature is not made, but born\u201d is not a value-neutral answer.<\/p>\n<p class=\"import-Normal\">Ancestry being the most important thing about a person became a popular idea twice in 20th century science. First, at the beginning of the century, when the <strong>eugenics<\/strong> movement in America called attention to \u201cfeeble-minded stocks,\u201d which usually referred to the poor or to immigrants (see Figure 17.4; and see Chapter 2). This movement culminated in Congress restricting the immigration of \u201cfeeble-minded races\u201d (said to include Jews and Italians) in 1924, and the Supreme Court declaring it acceptable for states to sterilize their \u201cfeeble-minded\u201d citizens involuntarily in 1927. After the Nazis picked up and embellished these ideas during World War II, Americans moved swiftly away from them in some contexts (e.g., for most people of European descent) while still strictly adhering in other contexts (e.g., Japanese internment camps and immigration restrictions).<\/p>\n<figure style=\"width: 374px\" class=\"wp-caption alignright\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image4-6.png\" alt=\"Historic photo. People sit in front of a structure with a \u201cEugenic and Health Exhibit&quot; banner.\" width=\"374\" height=\"262\" \/><figcaption class=\"wp-caption-text\">Figure 17.4: Eugenic and Health Exhibit, Fitter Families exhibit, and examination building, Kansas State Free Fair. Credit: <a href=\"https:\/\/www.dnalc.org\/view\/16328-Gallery-14-Eugenics-Exhibit-at-the-Kansas-State-Free-Fair-1920.html\">Gallery 14: Eugenics Exhibit at the Kansas State Free Fair, 1920 ID (ID 16328)<\/a> by <a href=\"https:\/\/www.dnalc.org\/\">Cold Spring Harbor<\/a> (Courtesy American Philosophical Society) is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/3.0\/us\/\">CC BY-NC-ND 3.0 License.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\">Ancestry again became paramount in the drumming up of public support for the Human Genome Project in the 1990s. Public support for sequencing the human genome was encouraged by a popular science campaign that featured books titled <em>The Book of Man <\/em>(Bodmer and McKie 1997), <em>The Human Blueprint <\/em>(Shapiro 1991), and <em>The Code of Codes<\/em> (Kevles and Hood 1993). These books generally promised cures for genetic diseases and a deeper understanding of the human condition. We can certainly identify progress in molecular genetics over the last couple of decades since the human genome was sequenced, but that progress has notably not been accompanied by cures for genetic diseases, nor by deeper understandings of the human condition.<\/p>\n<p class=\"import-Normal\">Even at the most detailed and refined levels of genetic analysis, we still don\u2019t have much of an understanding of the actual basis by which things seem to \u201crun in families.\u201d While the genetic basis of simple, if tragic, genetic diseases have become well-known\u2014such as sickle-cell anemia, cystic fibrosis, and Tay-Sachs\u2019 Disease\u2014we still haven\u2019t found the ostensible genetic basis for traits that are thought to have a strong genetic component. For example, a recent genetic summary found over 12,000 genetic sites that contributed to height yet still explained only about 40-50 percent of the variation in height among European ancestry but no more than 10-20 percent of variation of other ancestries, which we know strongly runs in families (Yengo et al. 2022).<\/p>\n<p class=\"import-Normal\">Partly in reaction to the reductionistic hype of the Human Genome Project, the study of epigenetics has become the subject of great interest. One famous natural experiment involves a Nazi-imposed famine in Holland over the winter of 1944\u20131945. Children born during and shortly after the famine experienced a higher incidence of certain health problems as adults, many decades later. Apparently, certain genes had been down-regulated early in development and remained that way throughout the course of life. Indeed, this modified regulation of the genes in response to the severe environmental conditions may have been passed on to their children.<\/p>\n<p class=\"import-Normal\">Obviously one\u2019s particular genetic constitution may play an important role in one\u2019s life trajectory. But overvaluing that role may have important social and political consequences. In the first place, genotypes are rendered meaningful in a cultural universe. Thus, if you live in a strongly patriarchal society and are born without a Y chromosome (since human males are chromosomally XY and females XX), your genotype will indeed have a strong effect upon your life course. So even though the variation is natural, the consequences are political. The mediating factors are the cultural ideas about how people of different sexes ought to be treated, and the role of the state in permitting certain people to develop and thrive. More broadly, there are implications for public education if variation in intelligence is genetic. There are implications for the legal system if criminality is genetic. There are implications for the justice system if sexual preference, or sexual identity, is genetic. There are implications for the development of sports talent if that is genetic. And yet, even for the human traits that are more straightforward to measure and known to be strongly heritable, the DNA base sequence variation seems to explain little.<\/p>\n<p class=\"import-Normal\">Genetic determinism or <strong>hereditarianism<\/strong> is the idea that \u201cthe creature is made, not born\u201d\u2014or, in a more recent formulation by James Watson, that \u201cour fate is in our genes.\u201d One of the major implications drawn from genetic determinism is that the feature in question must inevitably express itself; therefore, we can\u2019t do anything about it. Therefore, we might as well not fund the social programs designed to ameliorate economic inequality and improve people\u2019s lives, because their courses are fated genetically. And therefore, they don\u2019t deserve better lives.<\/p>\n<p class=\"import-Normal\">All of the \u201ctherefores\u201d in the preceding paragraph are open to debate. What is important is that the argument relies on a very narrow understanding of the role of genetics in human life, and it misdirects the causes of inequality from cultural to natural processes. By contrast, instead of focusing on genes and imagining them to place an invisible limit upon social progress, we can study the ways in which your DNA sequence does <em>not<\/em> limit your capability for self-improvement or fix your place in a social hierarchy. In general, two such avenues exist. First, we can examine the ways in which the human body responds and reacts to environmental variation: human adaptability and plasticity. This line of research began with the anthropometric studies of immigrants by Franz Boas in the early 20th century and has now expanded to incorporate the epigenetic inheritance of modified human DNA. And second, we can consider how human lives are shaped by social histories\u2014especially the structural inequalities within the societies in which they grow up.<\/p>\n<p class=\"import-Normal\">Although it arises and is refuted every generation, the radical hereditarian position (genetic determinism) perennially claims to speak for both science and evolution. It does not. It is the voice of a radical fringe\u2014perhaps naive, perhaps evil. It is not the authentic voice of science or of evolution. Indeed, keeping Charles Darwin\u2019s name unsullied by protecting it from association with bad science often seems like a full-time job. Culture and epigenetics are very much a part of the human condition, and their roles are significant parts of the complete story of human evolution.<\/p>\n<p><span style=\"background-color: #00ffff\"><span style=\"text-decoration: underline\">(Sterilization of Indigenous women in Canada)<\/span> (https:\/\/www.thecanadianencyclopedia.ca\/en\/article\/sterilization-of-indigenous-women-in-canada)\u00a0<\/span><\/p>\n<h2 class=\"import-Normal\">Adaptationism and the Panglossian Paradigm<\/h2>\n<p class=\"import-Normal\">The story of human evolution, and the evolution of all life for that matter, is never settled because evolution is ongoing. Additionally, because the conditions that shape evolutionary trajectories are not predetermined, evolution itself is emergent. Even during periods of ecological stability, when fewer macroevolutionary changes occur, populations of organisms continue to experience change. When ecological stability is disrupted, populations must adapt to the changes. Darwin explained in naturalistic terms how animals adapt to their environments: traits that contribute to an organism's ability to survive and reproduce in specific environments will become more common. The most \u201cfit\u201d\u2014those organisms best suited to the <em>current<\/em> environmental conditions in which they live\u2014have survived over eons of the history of life on earth to cocreate ecosystems full of animals and plants. Our own bodies are full of evident adaptations: eyes for seeing, ears for hearing, feet for walking on, and so forth.<\/p>\n<p class=\"import-Normal\">But what about hands? Feet are adapted to be primarily weight-bearing structures (rather than grasping structures, as in the apes) and that is what we primarily use them for. But we use our hands in many ways: for fine-scale manipulation, greeting, pointing, stimulating a sexual partner, writing, throwing, and cooking, among other uses. So which of these uses express what hands are \u201cfor,\u201d when all of them express what hands do?<\/p>\n<p class=\"import-Normal\">Gould and Lewontin (1979) illustrate the problem with assuming that the function of a trait defines its evolutionary cause. Consider the case of Dr. Pangloss\u2014the protagonistic of Voltaire\u2019s <em>Candide<\/em>\u2014who believed that we lived in the best of all possible worlds. Gould and Lewontin use his pronouncement that \u201cnoses were made for spectacles and so we have spectacles\u201d to demonstrate the problem with assuming any trait has evolved for a specific purpose. Identifying a function of a trait does not necessitate that the function is the ultimate cause of the trait. Individual traits are not under selection pressures in isolation; in fact, an entire organism must be able to survive and reproduce in their environment. When natural selection results in adaptations, changes that occur in some traits can have cascading effects throughout the phenotype and features that are not under selection pressure can also change.<\/p>\n<figure style=\"width: 279px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image3-5.png\" alt=\"Human hand is smaller with smaller fingers and smoother skin compared to a chimpanzee hand.\" width=\"279\" height=\"264\" \/><figcaption class=\"wp-caption-text\">Figure 17.5: Drawings of a human hand (left) and a chimpanzee hand (right). Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__\/\">Human and chimpanzee hand (Figure 2.16)<\/a> by Mary Nelson original to <a href=\"https:\/\/explorations.americananthro.org\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">There is an important lesson in recognizing that what things do in the present is not a good guide to understanding why they came to exist. Gunpowder was invented for entertainment\u2014only later was it adopted for killing people. The Internet was invented to decentralize computers in case of a nuclear attack\u2014and only later adopted for social media. Apes have short thumbs and use their hands in locomotion; our ancestors stopped using their hands in locomotion by about six million years ago and had fairly modern-looking hands by about two million years ago. We can speculate that a combination of selection for abstract thought and dexterity led to evolution of the human hand, with its capability for toolmaking that exceeds what apes can do (see Figure 17.5). But let\u2019s face it\u2014how many tools have you made today?<\/p>\n<p class=\"import-Normal\">Consequently, we are obliged to see the human foot as having a purpose to which it is adapted and the human hand as having multiple purposes, most of which are different from what it originally evolved for. Paleontologists Gould and Elisabeth Vrba suggested that an original use be regarded as an adaptation and any additional uses be called \u201c<strong>exaptations.<\/strong>\u201d Thus, we would consider the human hand to be an adaptation for toolmaking and an exaptation for writing. So how do we know whether any particular feature is an adaptation, like the walking foot, rather than an exaptation, like the writing hand? Or more broadly, how can we reason rigorously from what a feature does to what it evolved for?<\/p>\n<p class=\"import-Normal\">The answer to the question \u201cwhat did this feature evolve for?\u201d creates an origin myth. This origin myth contains three assumptions: (1) features can be isolated as evolutionary units; (2) there is a specific reason for the existence of any particular feature; and (3) there is a clear and simplistic explanation for why the feature evolved.<\/p>\n<figure style=\"width: 378px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image9-8.png\" alt=\"Head with images and human qualities drawn on it. Journal title printed at the bottom.\" width=\"378\" height=\"437\" \/><figcaption class=\"wp-caption-text\">Figure 17.6: According to the early 19th century science of phrenology, units of personality could be mapped onto units in the head, as shown on this cover of The Phrenology Journal. Credit: <a href=\"https:\/\/wellcomecollection.org\/works\/b6skynug\">Phrenology; Chart<\/a> [slide number 5278, photo number: L0000992, original print from Dr. E. Clark, The Phrenological Journal (Know Thyself)] by <a href=\"https:\/\/wellcomecollection.org\/\">Wellcome Collection<\/a>, is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/legalcode\">CC BY 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">The first assumption was appreciated a century ago as the \u201cunit-character problem.\u201d Are the units by which the body grows and evolves the same as units we name? This is clearly not the case: we have genes and we have noses, and we have genes that affect noses, but we don\u2019t have \u201cnose genes.\u201d What is the relationship between the evolving elements that we see, identify, and name, and the elements that biologically exist and evolve? It is hard to know, but we can use the history of science as a guide to see how that fallacy has been used by earlier generations. Back in the 19th century, the early anatomists argued that since the brain contained the mind, they could map different mental states (acquisitiveness, punctuality, sensitivity) onto parts of the brain. Someone who was very introspective, say, would have an enlarged introspection part of the brain, a cranial bulge to represent the hyperactivity of this mental state. The anatomical science was known as <strong>phrenology<\/strong>, and it was predicated on the false assumption that units of thought or personality or behavior could be mapped to distinct parts of the brain and physically observed (see Figure17.6). This is the fallacy of reification, imagining that something named is something real.<\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Long alt text: Side view of human head. At the top are the words \u201cKnow Thyself.\u201d On the upper head are small illustrations and word qualities such as \u201cfriendship,\u201d \u201cself-esteem,\u201d and \u201csecretiveness.\u201d On the lower part of the man\u2019s man\u2019s face are the words <em>The Phrenological Journal and Science of Health, A First Class Monthly<\/em>. The caption at the bottom reads: \u201cSpecially devoted to the \u2018.\u2019 Contains PHRENOLOGY and PHYSIOGNOMY, with all the SIGNS OF CHARACTER, and how to read them; ETHNOLOGY, or the Natural History of Man in all his relations.\u201d (All emphases in original.)<\/span><\/p>\n<figure style=\"width: 295px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image1-8-1.png\" alt=\"A black-and-white drawing of a chimpanzee head and face.\" width=\"295\" height=\"236\" \/><figcaption class=\"wp-caption-text\">Figure 17.7: Chimpanzees have big ears. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Chimpanzee_head_sketch.png\">Chimpanzee head sketch<\/a> by <a href=\"https:\/\/de.wikipedia.org\/wiki\/Benutzer:Roger_Zenner\">Roger Zenner<\/a>, original by Brehms Tierleben (1887), is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">The second assumption, that everything has a reason, has long been recognized as a core belief of religion. Our desire to impose order and simplicity on the workings of the universe, however, does not constrain it to obey simple and orderly causes. Magic, witchcraft, spirits, and divine agency are all powerful explanations for why things happen. Consequently, it is probably not a good idea to lump natural selection in with those. Sometimes things do happen for a reason, of course, but other times things happen as byproducts of other things, or for very complicated and entangled reasons, or for no reason at all. What phenomena have reasons and thereby merit explanation? Chimpanzees have very large testicles, and we think we know why: their promiscuous sexual behavior triggers intense competition for high sperm count. But chimpanzees also have very large ears, but much less scientific attention has been paid to this trait (see Figure 17.7). Why not? Why should there be a reason for chimp testicles but not for chimp ears? What determines the kinds of features that we try to explain, as opposed to the ones that we do not? Again, the assumption that any specific feature has a reason is metaphysical; that is to say, it may be true in any particular case, but to assume it in all cases is gratuitous.<\/p>\n<p class=\"import-Normal\">And third, the possibility of knowing what the reason for any particular feature is, assuming that it has one, is a challenge for evolutionary epistemology (the theory of how we know things). Consider the big adaptations of our lineage: bipedalism and language. Nobody doubts that they are good, and they evolved by natural selection, and we know how they work. But why did they evolve? If talking and walking are simply better than not talking and not walking, then why did they evolve in just a single branch of the ape lineage in the primate family tree? We don\u2019t know what bipedalism evolved for, although there are plenty of speculations: walking long distances, running long distances, cooling the head, seeing over tall grass, carrying babies, carrying food, wading, threatening, counting calories, sexual display, and so on. Neither do we know what language evolved for, although there are speculations: coordinating hunting, gossiping, manipulating others. But it is also possible that bipedality is simply the way that a small arboreal ape travels on the ground, if it isn\u2019t in the treetops. Or that language is simply the way that a primate with small canine teeth and certain mental propensities comes to communicate. If that were true, then there might be no reason for bipedality or language: having the unique suite of preconditions and a fortuitous set of circumstances simply set them in motion, and natural selection elaborated and explored their potentials. It is possible that walking and talking simply solved problems that no other lineage had ever solved; but even if so, the fact remains that the rest of the species in the history of life have done pretty well without having solved them.<\/p>\n<p class=\"import-Normal\">It is certainly very optimistic to think that all three assumptions (that organisms can be meaningfully atomized, that everything has a reason, and that we can know the reason) would be simultaneously in effect. Indeed, just as there are many ways of adapting (genetically, epigenetically, behaviorally, culturally), there are also many ways of being nonadaptive, which would imply that there is no reason at all for the feature in question.<\/p>\n<p class=\"import-Normal\">First, there is the element of randomness of population histories. There are more cases of sickle-cell anemia among sub-Saharan Africans than other peoples, and there is a reason for it: carriers of sickle-cell anemia have a resistance to malaria, which is more frequent in parts of Africa (as discussed in Chapters 4 and 14). But there are more cases of a blood disease called variegated porphyria, a rare genetic metabolic disorder, in the Afrikaners of South Africa (descendants of mostly Dutch settlers in the 17th century) than in other peoples, and there is no reason for it. Yet we know the cause: One of the founding Dutch colonial settlers had the <strong>allele<\/strong>\u2013a variant of a gene\u2013and everyone in South Africa with it today is her descendant. But that is not a reason\u2014that is simply an accident of history.<\/p>\n<p class=\"import-Normal\">Second, there is the potential mismatch between the past and the present. The value of a particular feature in the past may be changed as the environmental circumstances change. Our species is diurnal, and our ancestors were diurnal. But beginning around a few hundred thousand years ago, our ancestors could build fires, which extended the light period, which was subsequently further amplified by lamps and candles. And over the course of the 20th century, electrical power has made it possible for people to stay up very late when it is dark\u2014working, partying, worrying\u2014to a greater extent than any other closely related species. In other words, we evolved to be diurnal, yet we are now far more nocturnal than any of our recent ancestors or close relatives. Are we adapting to nocturnality? If so, why? Does it even make any sense to speak of the human occupation of a nocturnal ape niche, despite the fact that we empirically seem to be doing just that? And if so, does it make sense to ask what the reason for it is?<\/p>\n<p class=\"import-Normal\">Third, there is a genetic phenomenon known as a selective sweep, or the hitchhiker effect. Imagine three genes\u2014A, B, and C\u2014located very closely together on a chromosome. They each have several variants, or alleles, in the population. Now, for whatever reason, it becomes beneficial to have one of the B alleles, say B4; this B4 allele is now under strong positive selection. Obviously, we will expect future generations to be characterized by mostly B4. But what was B4 attached to? Because whatever A and C alleles were adjacent to it will also be quickly spread, simply by virtue of the selection for B4. Even if the A and C alleles are not very good, they will spread because of the good B4 allele between them. Eventually the linkage groups will break up because of genetic crossing-over in future generations. But in the meantime, some random version of genes A and C are proliferating in the species simply because they are joined to superior allele B4. And clearly, the A and C alleles are there because of selection\u2014but not because of selection <em>for<\/em> them!<\/p>\n<p class=\"import-Normal\">Fourth, some features are simply consequences of other properties rather than adaptations to external conditions. We already noted the phenomenon of allometric growth, in which some physical features have to outgrow others to maintain function at an increased size. Can we ask the reason for the massive brow ridges of <em>Homo erectus<\/em>, or are brow ridges simply what you get when you have a conjunction of thick skull bones, a large face, and a sloping forehead\u2014and, thus, again would have a cause but no reason?<\/p>\n<p class=\"import-Normal\">Fifth, some features may be underutilized and on the way out. What is the reason for our two outer toes? They aren\u2019t propulsive, they don\u2019t do anything, and sometimes they\u2019re just in the way. Obviously they are there because we are descended from ancestors with five digits on their hands and feet. Is it possible that a million years from now, we will just have our three largest toes, just as the ancestors of the horse lost their digits in favor of a single hoof per limb? Or will our outer toes find another use, such as stabilizing the landings in our personal jet-packs? For the time being, we can just recognize vestigiality as another nonadaptive explanation for the presence of a given feature.<\/p>\n<p class=\"import-Normal\">Finally, Darwin himself recognized that many obvious features do not help an animal survive. Some things may instead help an animal breed. The peacock\u2019s tail feathers do not help it eat, but they do help it mate. There is competition, but only against half of the species. Darwin called this <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1084\">sexual selection<\/a><\/strong>. Its result is not a fit to the environment but, rather, a fit to the opposite sex. In some species, that is literally the case, as the male and female genitalia have specific ways of anatomically fitting together. The specific form is less important than the specific match, so inquiring about the reason for a particular form of the reproductive anatomy may be misleading. The specific form may be effectively random, as long as it fits the opposite sex and is different from the anatomies of other species. Nor is sexual selection the only form of selection that can affect the body differently from natural selection. Competition might also take place between biological units other than organisms\u2014perhaps genes, perhaps cells, or populations, or species. The spread of cultural things, such as head-binding or cheap refined fructose or forced labor, can have significant effects upon bodies, which are also not adaptations produced by natural selection. They are often adaptive physiological responses to stresses but not the products of natural selection.<\/p>\n<p class=\"import-Normal\">With so many paths available by which a physical feature might have organically arisen without having been the object of natural selection, it is unwise to assume that any individual trait is an adaptation. And that generalization applies to the best-known, best-studied, and most materially based evolutionary adaptations of our lineage. But our cultural behaviors are also highly adaptive, so what about our most familiar social behaviors? Patriarchy, hierarchy, warfare\u2014are these adaptations? Do they have reasons? Are they good for something?<\/p>\n<p class=\"import-Normal\">This is where some sloppy thinking has been troublesome. What would it mean to say that patriarchy evolved by natural selection in the human species? If, on the one hand, it means that the human mind evolved by natural selection to be able to create and survive in many different kinds of social and political regimes, of which patriarchy is one, then biological anthropologists will readily agree. If, on the other hand, it means that patriarchy evolved by natural selection, that implies that patriarchy is genetically determined (since natural selection is a genetic process) and out-reproduced the alleles for other, more egalitarian, social forms. This in turn would imply that patriarchy is an adaptation and therefore of some beneficial value in the past and has become an ingrained part of human nature today. This would be bad news, say, if you harbored ambitions of dismantling it. Dismantling patriarchy in that case would be to go against nature, a futile gesture. In other words, this latter interpretation would be a naturalistic manifesto for a conservative political platform: don\u2019t try to dismantle the patriarchy, because it is within us, the product of evolution\u2014suck it up and live with it.<\/p>\n<p class=\"import-Normal\">Here, evolution is being used as a political instrument for transforming the human genome into an imaginary glass ceiling against equality. There is thus a convergence between the pseudo-biology of crude <strong>adaptationism <\/strong>(the idea that everything is the product of natural selection) and the pseudo-biology of hereditarianism. Naturalizing inequality is not the business of evolutionary theory, and it represents a difficult moral position for a scientist to adopt, as well as a poor scientific position.<\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ffff00\"><strong style=\"font-family: 'Cormorant Garamond', serif;font-size: 1.602em;background-color: #ffff00\">Evolution of the Anthropocene (to be reviewed)<\/strong><\/span><\/p>\n<figure style=\"width: 470px\" class=\"wp-caption alignright\"><img src=\"https:\/\/images.squarespace-cdn.com\/content\/v1\/5915c70e59cc6830f44a9d74\/1535724432724-FJ6FIAZI1B7RR65F3FMD\/ANTH_TFOS_DAN_02_16_SRC_WEB+%282%29.jpg\" alt=\"Dandora Landfill #3, Plastics Recycling, Nairobi, Kenya 2016\" width=\"470\" height=\"352\" \/><figcaption class=\"wp-caption-text\">comes from here https:\/\/www.edwardburtynsky.com\/projects\/the-anthropocene-project<\/figcaption><\/figure>\n<p>Under the previously explored Adaptationism and Panglossian Paradigm, it is explained that human evolution is constantly occurring even throughout periods of ecological stability. While this acknowledges evolution as an ongoing process of change, it fails to explore the implications of such on the alteration of other species and ecosystems.<\/p>\n<p>The emergence of the Anthropocene, driven by human activity, though not recognized as an official epoch, is seen as a transformative event comparable to other major historical shifts such as the Ordovician Biodiversification (UNESCO, 2024). Given its scale, it is crucial to inform scholars about the impact of our social and cultural evolution on the rest of the world. Richard Robbins\u2019 Global Problems and Culture of Capitalism explains how the modern culture of consumption has been extremely successful at accommodating populations of people far larger than previously possible. Robbins claims that the globalization attributed to capitalism has allowed the world to make full use of its environmental resources, providing necessities and innovative technologies to humans all over the world (Robbins &amp; Dowty, 2019). In other words, capitalism is an anthropocentric cultural system that highly benefits humans and facilitates our survival with little regard to the development and survival of other forms of life. It would be highly relevant to introduce the idea that our cultural evolution and capacity to modify the environment to meet our needs have established new environmental conditions in which the human species' survival and reproduction rate expand at the detriment of ecosystems and endangerment of other primates and non-human species.<\/p>\n<p>According to the International Union for Conservation of Nature\u2019s Red List of Threatened Species, there are currently over 169,000 species listed, with more than 47,000 species at risk of extinction \u2014 including 41% of amphibians, 26% of mammals, 26% of freshwater fishes, 12% of birds, and many others (IUCN, 2025). Human lifestyles are causing changes that\u2014if not taken into consideration\u2014could lead to our extinction as a species. The recognition that our evolutionary behavioural development is causing environmental destruction may be the first step for our species to take accountability for the damage that it is causing to others and prevent further damage.<\/p>\n<p><span style=\"background-color: #ff99cc;font-family: 'Cormorant Garamond', serif;font-size: 1.602em;font-weight: bold\">Concluding Thoughts<\/span><\/p>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Now that you have finished reading this chapter, you are equipped to understand the historical and political dimensions of evolution. Evolution is an ongoing process of change and diversification. Evolutionary theory is a tool that we use to understand this process. The development of evolutionary theory is shaped both by scientific innovation and political engagement. Since Darwin first articulated natural selection as an observable mechanism by which species adapt to their environments, our understanding of evolution has grown. Initially, scientists focused on the adaptive aspects of evolution. However, with the emergence of genetics, our understanding of heredity and the level at which evolution acts has changed. Genetics led to a focus on the molecular dimensions of evolution. For some, this focus resulted in reductive accounts of evolution. Further developments in our understanding of evolution shifted our view to epigenetic processes and how organisms shape their own evolutionary pressures (e.g., niche construction).<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Evolutionary theory will continue to develop in the future as we invent new technologies, describe new dimensions of biology, and experience cultural changes. Current innovations in evolutionary theory are asking us to consider evolutionary forces beyond natural selection and genetics to include the ways organisms shape their environments (niche construction), inheritances beyond genetics (inclusive inheritance), constraints on evolutionary change (developmental bias), and the ability of bodies to change in response to external factors (plasticity). The future of evolutionary theory looks bright as we continue to explore these and other dimensions. Biological anthropology is well-positioned to be a lively part of this conversation, as it extends standard evolutionary theory by considering the role of culture, social learning, and human intentionality in shaping the evolutionary trajectories of humans (Zeder 2018). Remember, at root, human evolutionary theory consists of two propositions: (1) the human species is descended from other similar species and (2) natural selection has been the primary agent of biological adaptation. Pretty much everything else is subject to some degree of contestation.<\/span><\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Review Questions<\/h2>\n<ul>\n<li class=\"import-Normal\">How is the study of your ancestors biopolitical, not just biological? Does that make it less scientific or differently scientific?<\/li>\n<li class=\"import-Normal\">What was gained by reducing organisms to genotypes and species to gene pools? What is gained by reintroducing bodies and species into evolutionary studies?<\/li>\n<li class=\"import-Normal\">How do genetic or molecular studies complement anatomical studies of evolution?<\/li>\n<li class=\"import-Normal\">How are you reducible to your ancestry? If you could meet your ancestors from the year 1700 (and you would have well over a thousand of them!), would their lives be meaningfully similar to yours? Would you even be able to communicate with them?<\/li>\n<li class=\"import-Normal\">The molecular biologist Fran\u00e7ois Jacob argued that evolution is more like a tinkerer than an engineer. In what ways do we seem like precisely engineered machinery, and in what ways do we seem like jerry-rigged or improvised contraptions?<\/li>\n<li class=\"import-Normal\">How might biological anthropology contribute to future developments in evolutionary theory?<\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\">Key Terms<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\"><strong>Adaptation<\/strong>: A fit between the organism and environment.<\/p>\n<p class=\"import-Normal\"><strong>Adaptationism<\/strong>: The idea that everything is the product of natural selection.<\/p>\n<p class=\"import-Normal\"><strong>Allele<\/strong>: A genetic variant.<\/p>\n<p class=\"import-Normal\"><strong>Allometry<\/strong>: The differential growth of body parts.<\/p>\n<p class=\"import-Normal\"><strong>Canalization<\/strong>: The tendency of a growing organism to be buffered toward normal development.<\/p>\n<p class=\"import-Normal\"><strong>Epigenetics<\/strong>: The study of how genetically identical cells and organisms (with the same DNA base sequence) can nevertheless differ in stably inherited ways.<\/p>\n<p class=\"import-Normal\"><strong>Eugenics<\/strong>: An idea that was popular in the 1920s that society should be improved by breeding \u201cbetter\u201d kinds of people.<\/p>\n<p class=\"import-Normal\"><strong>Evo-devo<\/strong>: The study of the origin of form; a contraction of \u201cevolutionary developmental biology.\u201d<\/p>\n<p class=\"import-Normal\"><strong>Exaptation<\/strong>: An additional beneficial use for a biological feature.<\/p>\n<p class=\"import-Normal\"><strong>Extinction<\/strong>: The loss of a species from the face of the earth.<\/p>\n<p class=\"import-Normal\"><strong>Gene<\/strong>: A stretch of DNA with an identifiable function (sometimes broadened to include any DNA with recognizable structural features as well).<\/p>\n<p class=\"import-Normal\"><strong>Gene pool<\/strong>: Hypothetical summation of the entire genetic composition of population or species.<\/p>\n<p class=\"import-Normal\"><strong>Genotype<\/strong>: Genetic constitution of an individual organism.<\/p>\n<p class=\"import-Normal\"><strong>Hereditarianism<\/strong>: The idea that genes or ancestry is the most crucial or salient element in a human life. Generally associated with an argument for natural inequality on pseudo-genetic grounds.<\/p>\n<p class=\"import-Normal\"><strong>Hox genes<\/strong>: A group of related genes that control for the body plan of an embryo along the head-tail axis.<\/p>\n<p class=\"import-Normal\"><strong>Inheritance of acquired characteristics<\/strong>: The idea that you pass on the features that developed during your lifetime, not just your genes; also known as Lamarckian inheritance.<\/p>\n<p class=\"import-Normal\"><strong>Natural selection<\/strong>: A consistent bias in survival and fertility, leading to the overrepresentation of certain features in future generations and an improved fit between an average member of the population and the environment.<\/p>\n<p class=\"import-Normal\"><strong>Niche construction<\/strong>: The active engagement by which species transform their surroundings in favorable ways, rather than just passively inhabiting them.<\/p>\n<p class=\"import-Normal\"><strong>Phenotype<\/strong>: Observable manifestation of a genetic constitution, expressed in a particular set of circumstances. The suite of traits of an organism.<\/p>\n<p class=\"import-Normal\"><strong>Phrenology<\/strong>: The 19th-century anatomical study of bumps on the head as an indication of personality and mental abilities.<\/p>\n<p class=\"import-Normal\"><strong>Plasticity<\/strong>: The tendency of a growing organism to react developmentally to its particular conditions of life.<\/p>\n<p class=\"import-Normal\"><strong>Punctuated equilibria<\/strong>: The idea that species are stable through time and are formed very rapidly relative to their duration. (The opposite theory, that species are unstable and constantly changing through time, is called phyletic gradualism.)<\/p>\n<p class=\"import-Normal\"><strong>Scientific racism<\/strong>: The use of pseudoscientific evidence to support or legitimize racial hierarchy and inequality.<\/p>\n<p class=\"import-Normal\"><strong>Sexual selection<\/strong>: Natural selection arising through preference by one sex for certain characteristics in individuals of the other sex.<\/p>\n<p class=\"import-Normal\"><strong>Species selection<\/strong>: A postulated evolutionary process in which selection acts on an entire species population, rather than individuals.<\/p>\n<h2 class=\"import-Normal\">About the Authors<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image2-4-1.jpg\" alt=\"A bearded man wearing glasses smiles at the camera. \" width=\"202\" height=\"218\" \/><\/p>\n<h3 class=\"import-Normal\"><strong>Jonathan Marks, Ph.D.<\/strong><\/h3>\n<p class=\"import-Normal\">University of North Carolina at Charlotte, <a class=\"rId41\" href=\"mailto:jmarks@uncc.edu\">jmarks@uncc.edu<\/a><\/p>\n<p class=\"import-Normal\">Jonathan Marks is Professor of Anthropology at the University of North Carolina at Charlotte. He has published many books and articles on broad aspects of biological anthropology. In 2006 he was elected a Fellow of the American Association for the Advancement of Science. In 2012 he was awarded the First Citizen\u2019s Bank Scholar\u2019s Medal from UNC Charlotte. In recent years he has been a Visiting Research Fellow at the ESRC Genomics Forum in Edinburgh, a Visiting Research Fellow at the Max Planck Institute for the History of Science in Berlin, and a Templeton Fellow at the Institute for Advanced Study at Notre Dame. His work has received the W. W. Howells Book Prize and the General Anthropology Division Prize for Exemplary Cross-Field Scholarship from the American Anthropological Association as well as the J. I. Staley Prize from the School for Advanced Research. Two of his books are titled <em>What It Means to Be 98% Chimpanzee<\/em> and <em>Why I Am Not a Scientist<\/em>, but actually he is about 98 percent scientist and not a chimpanzee.<\/p>\n<p class=\"import-Normal\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image10-3.jpg\" alt=\"A bearded man wearing a fedora hat looks off in the distance. \" width=\"232\" height=\"232\" \/><\/p>\n<h3 class=\"import-Normal\"><strong>Adam P. Johnson, M.A.<\/strong><\/h3>\n<p class=\"import-Normal\">University of North Carolina at Charlotte\/University of Texas at San Antonio, <a class=\"rId43\" href=\"mailto:ajohn344@uncc.edu\">ajohn344@uncc.edu<\/a><\/p>\n<p class=\"import-Normal\">Adam Johnson is a doctoral candidate at the University of Texas at San Antonio and part-time lecturer at the University of North Carolina at Charlotte. He earned his M.A. in anthropology at UNC-Charlotte in 2017 and will complete his Ph.D. in anthropology at UTSA by 2024. His interests include human-animal relations, science studies, primate behavior, ecology, and the history of anthropology. His recent research project analyzes the social, historical, political, and evolutionary dimensions that shape human-javelina encounters. His goal is to understand how humans and animals find ways to get along in a precarious world.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration <strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\">Ackermann, Rebecca Rogers, Alex Mackay, and Michael L. Arnold. 2016. \u201cThe Hybrid Origin of \u2018Modern\u2019 Humans.\u201d <em>Evolutionary Biology<\/em> 43 (1): 1\u201311.<\/p>\n<p class=\"import-Normal\">Bateson, Patrick, and Peter Gluckman. 2011. <em>Plasticity, Robustness, Development and Evolution<\/em>. New York: Cambridge University Press.<\/p>\n<p class=\"import-Normal\">Cosans, Christopher E. 2009. <em>Owen's Ape and Darwin's Bulldog: Beyond Darwinism and Creationism<\/em>. Bloomington, IN: Indiana University Press.<\/p>\n<p class=\"import-Normal\">Desmond, Adrian, and James Moore. 2009. <em>Darwin's Sacred Cause: How a Hatred of Slavery Shaped Darwin's Views on Human Evolution<\/em>. New York: Houghton Mifflin Harcourt.<\/p>\n<p class=\"import-Normal\">Dobzhansky, Theodosius, Francisco J. Ayala, G. Ledyard Stebbins, and James W. Valentine. 1977. <em>Evolution<\/em>. San Francisco: W.H. Freeman and Company.<\/p>\n<p class=\"import-Normal\">Fuentes, Agust\u00edn. 2017. <em>The Creative Spark: How Imagination Made Humans Exceptional<\/em>. New York: Dutton.<\/p>\n<p class=\"import-Normal\">Gould, Stephen J. 2003.<em> The Structure of Evolutionary Theory<\/em>. Cambridge, MA: Harvard University Press.<\/p>\n<p class=\"import-Normal\">Haraway, Donna J. 1989. <em>Primate Visions: Gender, Race, and Nature in the World of Modern Science<\/em>. New York: Routledge.<\/p>\n<p class=\"import-Normal\">Huxley, Thomas. 1863. <em>Evidence as to Man's Place in Nature<\/em>. London: Williams &amp; Norgate.<\/p>\n<p class=\"import-Normal\">Jablonka, Eva, and Marion J. Lamb. 2005. <em>Evolution in Four Dimensions: Genetic, Epigenetic, Behavioral, and Symbolic Variation in the History of Life<\/em>. Cambridge, MA: The MIT Press.<\/p>\n<p class=\"import-Normal\">Kuklick, Henrika, ed. 2008. <em>A New History of Anthropology<\/em>. New York: Blackwell.<\/p>\n<p class=\"import-Normal\">Laland, Kevin N., Tobias Uller, Marcus W. Feldman, Kim Sterelny, Gerd B. Muller, Armin Moczek, Eva Jablonka, and John Odling-Smee. 2015. \u201cThe Extended Evolutionary Synthesis: Its Structure, Assumptions and Predictions.\u201d <em>Proceedings of the Royal Society, Series B<\/em> 282 (1813): 20151019.<\/p>\n<p class=\"import-Normal\">Lamarck, Jean Baptiste. 1809. <em>Philosophie Zoologique<\/em>. Paris: Dentu.<\/p>\n<p class=\"import-Normal\">Landau, Misia. 1991. <em>Narratives of Human Evolution<\/em>. New Haven: Yale University Press.<\/p>\n<p class=\"import-Normal\">Lee, Sang-Hee. 2017. <em>Close Encounters with Humankind: A Paleoanthropologist Investigates Our Evolving Species<\/em>. New York: W. W. Norton.<\/p>\n<p class=\"import-Normal\">Livingstone, David N. 2008. <em>Adam's Ancestors: Race, Religion, and the Politics of Human Origins<\/em>. Baltimore: Johns Hopkins University Press.<\/p>\n<p class=\"import-Normal\">Marks, Jonathan. 2015. <em>Tales of the Ex-Apes: How We Think about Human Evolution<\/em>. Berkeley, CA: University of California Press.<\/p>\n<p class=\"import-Normal\">Pigliucci, Massimo. 2009. \u201cThe Year in Evolutionary Biology 2009: An Extended Synthesis for Evolutionary Biology.\u201d <em>Annals of the New York Academy of Sciences<\/em> 1168: 218\u2013228.<\/p>\n<p class=\"import-Normal\">Simpson, George Gaylord. 1949. <em>The Meaning of Evolution: A Study of the History of Life and of Its Significance for Man<\/em>. New Haven: Yale University Press.<\/p>\n<p class=\"import-Normal\">Sommer, Marianne. 2016.<em> History Within: The Science, Culture, and Politics of Bones, Organisms, and Molecules<\/em>. Chicago: University of Chicago Press.<\/p>\n<p class=\"import-Normal\">Stoczkowski, Wiktor. 2002. <em>Explaining Human Origins: Myth, Imagination and Conjecture<\/em>. New York: Cambridge University Press.<\/p>\n<p class=\"import-Normal\">Tattersall, Ian, and Rob DeSalle. 2019. <em>The Accidental Homo sapiens: Genetics, Behavior, and Free Will<\/em>. New York: Pegasus.<\/p>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\">Barton, Robert A. 1996. \"Neocortex Size and Behavioural Ecology in Primates.\" <em>Proceedings of the Royal Society of London. Series B: Biological Sciences<\/em> 263 (1367): 173\u2013177.<\/p>\n<p class=\"import-Normal\">Bodmer, Walter, and Robin McKie. 1997. <em>The Book of Man: The Hman Genome Project and the Quest to Discover our Genetic Heritage.<\/em> Oxford University Press.<\/p>\n<p class=\"import-Normal\">Darwin, Charles. 1859.<em> On the Origin of Species by Means of Natural Selection, or, the Preservation of Favoured Races in the Struggle for Life<\/em>. London: J. Murray.<\/p>\n<p class=\"import-Normal\">Darwin, Charles. 1871. <em>The Descent of Man, and Selection in Relation to Sex.<\/em> London: J. Murray.<\/p>\n<p class=\"import-Normal\">Dawkins, Richard. 1976. <em>The Selfish Gene. <\/em>Oxford University Press.<\/p>\n<p class=\"import-Normal\">Deacon, T. W. 1998. <em>The Symbolic Species: The Co-evolution of Language and the Brain<\/em>. W. W. Norton &amp; Company.<\/p>\n<p class=\"import-Normal\">Eldredge, N., and S. J. Gould. 1972. \"Punctuated Equilibria: An Alternative to Phyletic Gradualism.\" In <em>Models in Paleobiology<\/em>, edited by T. J. Schopf, 82\u2013115. San Francisco: W. H. Freeman.<\/p>\n<p class=\"import-Normal\">Gould, Stephen J. 2003.<em> The Structure of Evolutionary Theory<\/em>. Cambridge, MA: Harvard University Press.<\/p>\n<p class=\"import-Normal\">Gould, Stephen J. 1996. <em>Mismeasure of Man<\/em>. New York: WW Norton &amp; Company.<\/p>\n<p class=\"import-Normal\">Gould, Stephen Jay, and Richard C. Lewontin. 1979. \"The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme.\" <em>Proceedings of the Royal Society of London. Series B: Biological Sciences<\/em> 205 (1151): 581\u2013598.<\/p>\n<p class=\"import-Normal\">Haeckel, Ernst. 1868. <em>Nat\u00fcrliche Sch\u00f6pfungsgeschichte<\/em>. Berlin: Reimer.<\/p>\n<p class=\"import-Normal\">Huxley, Thomas Henry. 1863. <em>Evidence as to Man\u2019s Place in Nature. <\/em>London: Williams and Norgate.<\/p>\n<p class=\"import-Normal\">Kaufman, Thomas C., Mark A. Seeger, and Gary Olsen. 1990. \"Molecular and Genetic Organization of the Antennapedia Gene Complex of <em>Drosophila melanogaster<\/em>.\" <em>Advances in Genetics<\/em> 27: 309\u2013362.<\/p>\n<p class=\"import-Normal\">Kellogg, Vernon. 1917. <em>Headquarters Nights<\/em>. Boston: The Atlantic Monthly Press.<\/p>\n<p class=\"import-Normal\">Kevles, Daniel J., and Leroy Hood. 1993. <em>The Code of Codes: Scientific and Social Issues in the Human Genome Project<\/em>. Cambridge, MA: Harvard University Press.<\/p>\n<p class=\"import-Normal\">Lewontin, Richard, Steven Rose, and Leon Kamin. 2017. <em>Not in Our Genes\u202f: Biology, Ideology, and Human Nature<\/em>, 2nd ed. Chicago: Haymarket Books.<\/p>\n<p class=\"import-Normal\">Lloyd, Elisabeth A., and Stephen J. Gould. 1993. \"Species Selection on Variability.\" <em>Proceedings of the National Academy of Sciences<\/em> 90 (2): 595\u2013599.<\/p>\n<p class=\"import-Normal\">Marks, Jonathan. 2015. \u201cThe Biological Myth of Human Evolution.\u201d In <em>Biologising the Social Sciences: Challenging Darwinian and Neuroscience Explanations<\/em>, edited by David Canter and David A. Turner, 59\u201378. London: Routledge.<\/p>\n<p class=\"import-Normal\">Monypenny, William Flavelle, and George Earle Buckle. 1929. <em>The Life of Benjamin Disraeli, Earl of Beaconsfield, Volume II: 1860\u20131881<\/em>. London: John Murray.<\/p>\n<p class=\"import-Normal\">Potts, Rick. 1998. \u201cVariability Selection in Hominid Evolution.\u201d <em>Evolutionary Anthropology <\/em><em>7<\/em><em>:<\/em> 81\u201396.<\/p>\n<p class=\"import-Normal\">Punnett, R. C. 1905. <em>Mendelism<\/em>. Cambridge: Macmillan and Bowes.<\/p>\n<p class=\"import-Normal\">Shapiro, Robert. 1991. <em>The Human Blueprint: The Race to Unlock the Secrets of Our Genetic Script.<\/em> New York: St. Martin\u2019s Press.<\/p>\n<p class=\"import-Normal\">Shultz, Susanne, Emma Nelson, and Robin Dunbar. 2012. \"Hominin Cognitive Evolution: Identifying Patterns and Processes in the Fossil and Archaeological Record.\" <em>Philosophical Transactions of the Royal Society B: Biological Sciences<\/em> 367 (1599): 2130\u20132140.<\/p>\n<p class=\"import-Normal\">Spencer, Herbert. 1864. <em>Principles of Biology.<\/em> London: Williams and Norgate.<\/p>\n<p class=\"import-Normal\">Watson, James D. 1990. \"The Human Genome Project: Past, Present, and Future.\" <em>Science<\/em> 248 (4951): 44\u201349.<\/p>\n<p class=\"import-Normal\">Yengo, L., Vedantam, S., Marouli, E., Sidorenko, J., Bartell, E., Sakaue, S., Graff, M., Eliasen, A.U., Jiang, Y., Raghavan, S. and Miao, J., 2022. A saturated map of common genetic variants associated with human height. <em>Nature<\/em>, <em>610 <\/em>(7933): 704-712.<\/p>\n<p class=\"import-Normal\">Zeder, Melinda A. 2018. \"Why Evolutionary Biology Needs Anthropology: Evaluating Core Assumptions of the Extended Evolutionary Synthesis.\" <em>Evolutionary Anthropology: Issues, News, and Reviews<\/em> 27 (6): 267\u2013284.<\/p>\n<\/div>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1792\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1792\"><div tabindex=\"-1\"><\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1778\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1778\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\">Ashley Kendell, Ph.D., California State University, Chico<\/p>\n<p class=\"import-Normal\">Alex Perrone, M.A., M.S.N, R.N., P.H.N., Butte Community College<\/p>\n<p class=\"import-Normal\">Colleen Milligan, Ph.D., California State University, Chico<\/p>\n<p class=\"import-Normal\"><em>This chapter is a revision from \"<\/em><a class=\"rId7\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\"><em>Chapter 15: Bioarchaeology and Forensic Anthropology<\/em><\/a><em>\u201d by Ashley Kendell, Alex Peronne, and Colleen Milligan. In <\/em><a class=\"rId8\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, first edition<\/em><\/a><em>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under <\/em><a class=\"rId9\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\"><em>CC BY-NC 4.0<\/em><\/a><em>. <\/em><\/p>\n<p class=\"import-Normal\"><strong>Content Warning and Disclaimer:<\/strong> This chapter includes images of human remains as well as discussions centered on human skeletal analyses. All images are derived from casts, sketches, nonhuman skeletal material, as well as non-Indigenous skeletal materials curated within the CSU, Chico Human Identification Lab, and the Hartnett-Fulginiti donated skeletal collection.<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\"><span style=\"color: #ffffff\">Learning Objectives<\/span><\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Define forensic anthropology as a subfield of biological anthropology.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Describe the seven steps carried out during skeletal analysis.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Outline the four major components of the biological profile.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Contrast the four categories of trauma.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Explain how to identify the different taphonomic agents that alter bone.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Discuss ethical considerations for forensic anthropology.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1214\">Forensic anthropology<\/a><\/strong> is a subfield of biological anthropology and an applied area of anthropology. Forensic anthropologists use skeletal analysis to gain information about humans in the present or recent past, then they apply this information within a medicolegal context. This means that forensic anthropologists specifically conduct their analysis on recently deceased individuals (typically within the last 50 years) as part of investigations by law enforcement. Forensic anthropologists can assist law enforcement agencies in several different ways, including aiding in the identification of human remains whether they are complete, fragmentary, burned, scattered, or decomposed. Additionally, forensic anthropologists can help determine what happened to the deceased at or around the time of death as well as what processes acted on the body after death (e.g., whether the remains were scattered by animals, whether they were buried in the ground, or whether they remained on the surface as the soft tissue decomposed).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Many times, because of their expertise in identifying human skeletal remains, forensic anthropologists are called to help with outdoor search-and-recovery efforts, such as locating remains scattered across the surface or carefully excavating and documenting buried remains. In other cases, forensic anthropologists recover remains after natural disasters or accidents, such as fire scenes, and can help identify whether each bone belongs to a human or an animal. Forensic anthropology spans a wide scope of contexts involving the law, including incidences of mass disasters, genocide, and war crimes.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">A point that can be somewhat confusing for students is that although the term <em>forensic<\/em> is included in this subfield of biological anthropology, there are many forensic techniques that are not included in the subfield. Almost exclusively, forensic anthropology deals with skeletal analysis. While this can include the comparison of antemortem (before death) and postmortem (after death) radiographs to identify whether remains belong to a specific person, or using photographic superimposition of the cranium, it does not include analyses beyond the skeleton. For example, blood-spatter analysis, DNA analysis, fingerprints, and material evidence collection do not fall under the scope of forensic anthropology.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">So, what can forensic anthropologists glean from bones alone? Forensic anthropologists can address a number of questions about a human individual based on their skeletal remains. Some of those questions are as follows: How old was the person? Was the person biologically male or female? How tall was the person? What happened to the person at or around their time of death? Were they sick? The information from the skeletal analysis can then be matched with missing persons records, medical records, or dental records, aiding law enforcement agencies with identifications and investigations.<\/p>\n<h2 class=\"import-Normal\">Skeletal Analysis<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropology relies on skeletal analysis to reveal information about the deceased. <span style=\"background-color: #00ffff\">The methodology and approaches outlined below are specific to the United States.<\/span> Forensic anthropological methods differ depending on the country conducting an investigation. In the United States, there are typically seven steps or questions to the process:<\/p>\n<ul>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Is it bone?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Is it human?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Is it modern or archeological?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">How many individuals are present or what is the minimum number of individuals (MNI)?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Who is it?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Is there evidence of trauma before or around the time of death?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">What happened to the remains after death?<\/li>\n<\/ul>\n<h3 class=\"import-Normal\"><strong>Is It Bone?<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">One of the most important steps in any skeletal analysis starts with determining whether or not material suspected to be bone is in fact bone. Though it goes without saying that a forensic anthropologist would only carry out analysis on bone, this step is not always straightforward. Whole bones are relatively easy to identify, but determining whether or not something is bone becomes more challenging once it becomes fragmentary. As an example, in high heat such as that seen on fire scenes, bone can break into pieces. During a house fire with fatalities, firefighters watered down the burning home. After the fire was extinguished, the sheetrock (used to construct the walls of the home) was drenched and crumbled. The crumbled sheetrock was similar in color and form to burned, fragmented bone, therefore mistakable for human remains (Figure 15.1). Forensic anthropologists on scene were able to separate the bones from the construction material, helping to confirm the presence of bone and hence the presence of individual victims of the fire. In this case, forensic anthropologists were able to recognize the anatomical and layered structure of bone and were able to distinguish it from the uniform and unlayered structure of sheetrock.<\/p>\n<p class=\"import-Normal\"><strong><img class=\"aligncenter\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/06\/image3.png\" alt=\"Long rectangular sheetrock with exposed porous surface.\" width=\"182\" height=\"208\" \/><\/strong><\/p>\n<figure style=\"width: 372px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image6-1.png\" alt=\"Two examples of sheetrock with dried or burnt surfaces.\" width=\"372\" height=\"210\" \/><figcaption class=\"wp-caption-text\">Figure 15.1: Burned sheetrock used as building material appears similar to human bone but can be differentiated by the fact that it is the same density throughout. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Example of burned sheetrock (Figure 15.1)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">As demonstrated by the example above, both the macrostructure (visible with the naked eye) and microstructure (visible with a microscope) of bone are helpful in bone identification. Bones are organs in the body made up of connective tissue. The connective tissue is hardened by a mineral deposition, which is why bone is rigid in comparison to other connective tissues such as cartilage (Tersigni-Tarrant and Langley 2017, 82\u201383; White and Folkens 2005, 31). In a living body, the mineralized tissue does not make up the only component of bone\u2014there are also blood, bone marrow, cartilage, and other types of tissues. However, in dry bone, two distinct layers of the bone are the most helpful for identification. The outer layer is made up of densely arranged osseous (bone) tissue called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1216\">compact (cortical) bone<\/a><\/strong>. The inner layer is composed of much more loosely organized, porous bone tissue whose appearance resembles that of a sponge, hence the name <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1218\"><strong>spongy (trabecular) bone<\/strong><\/a>. Knowing that most bone contains both layers helps with the macroscopic identification of bone (Figures 15.2, 15.3). For example, a piece of coconut shell might look a lot like a fragment of a human skull bone. However, closer inspection will demonstrate that coconut shell only has one very dense layer, while bone has both the compact and spongy layers.<\/p>\n<figure style=\"width: 380px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image27-1.png\" alt=\"Drawing showing thick exterior compact bone and porous internal cortical bone.\" width=\"380\" height=\"371\" \/><figcaption class=\"wp-caption-text\">Figure 15.2: Cross section of human long bone with compact and cortical bone layers visible. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Cross section of human long bone (Figure 15.2)<\/a> original to<a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"> Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p>&nbsp;<\/p>\n<figure style=\"width: 364px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image25-2.png\" alt=\"Cranial bone cross section called a periosteum with spongy bone (diploe) and compact bone labeled. Compact bone is a thin slice at the top and bottom and is smooth and hard. Spongy bone is in the middle and has irregular holes and indentations throughout. \" width=\"364\" height=\"184\" \/><figcaption class=\"wp-caption-text\">Figure 15.3: Cranial anatomy is slightly different as compared to that of a long bone in cross section. The compact (cortical) bone layers sandwich the spongy (trabecular) bone. One layer of compact bone forms the very outer surface of the skull and the other lines the internal surface of the skull. Credit: <a href=\"https:\/\/cnx.org\/contents\/FPtK1zmh@6.27:kwbeYj9S@3\/Bone-Structure\">Anatomy of a Flat Bone (Anatomy &amp; Physiology, Figure 6.3.3)<\/a> by<a href=\"https:\/\/openstax.org\/\"> OpenStax<\/a> is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/\"> CC BY 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The microscopic identification of bone relies on knowledge of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1220\">osteons<\/a><\/strong>, or bone cells (Figure 15.4). Under magnification, bone cells are visible in the outer, compact layer of bone. The bone cells are arranged in a concentric pattern around blood vessels for blood supply. The specific shape of the cells can help differentiate, for example, a small piece of PVC (white plastic) pipe from a human bone fragment (Figure 15.5).<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 340px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image14-3.png\" alt=\"Microscope image showing clustered osteons. Each has many rings and a dark center.\" width=\"340\" height=\"218\" \/><figcaption class=\"wp-caption-text\">Figure 15.4: Bone microstructure (osteons). Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Bone_(248_12)_Bone_cross_section.jpg\">Bone (248 12) Bone cross section<\/a> by <a href=\"https:\/\/cs.wikipedia.org\/wiki\/Josef_Reischig\">Doc. RNDr. Josef Reischig, CSc.<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p>&nbsp;<\/p>\n<figure style=\"width: 332px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image7-1.png\" alt=\"Flat, white section of PVC. Edges are broken and surface rough.\" width=\"332\" height=\"268\" \/><figcaption class=\"wp-caption-text\">Figure 15.5: Fragments of plastic PVC pipe, such as those seen in this photo, may be mistaken for human bone. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Example of PVC pipe<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Is It Human?<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Once it has been determined that an object is bone, the next logical step is to identify whether the bone belongs to a human or an animal. Forensic anthropologists are faced with this question in everyday practice because human versus nonhuman bone identification is one of the most frequent requests they receive from law enforcement agencies.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">There are many different ways to distinguish human versus nonhuman bone. The morphology (the shape\/form) of human bone is a good place for students to start. Identifying the 206 bones in the adult human skeleton and each bone\u2019s distinguishing features (muscle attachment sites, openings and grooves for nerves and blood vessels, etc.) is fundamental to skeletal analysis.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Nevertheless, there are many animal bones and human bones that look similar. For example, the declawed skeleton of a bear paw looks a lot like a human hand, pig molars appear similar to human molars, and some smaller animal bones might be mistaken for those of an infant. To add to the confusion, fragmentary bone may be even more difficult to identify as human or nonhuman. However, several major differences between human and nonhuman vertebrate bone help distinguish the two.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropologists pay special attention to the density of the outer, compact layer of bone in both the cranium and in the long bones. Human cranial bone has three distinctive layers. The spongy bone is sandwiched between the outer (ectocranial) and inner (endocranial) compact layers. In most other mammals, the distinction between the spongy and compact layers is not always so definite. Secondly, the compact layer in nonhuman mammal long bones can be much thicker than observed in human bone. Due to the increased density of the compact layer, nonhuman bone tends to be heavier than human bone (Figure 15.6).<\/p>\n<figure style=\"width: 399px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image11-2.png\" alt=\"Ring-like cross section of bone.\" width=\"399\" height=\"266\" \/><figcaption class=\"wp-caption-text\">Figure 15.6: The compact layer of this animal bone is very thick, with almost no spongy bone visible. Compare with Figure 15.2 to visualize the difference in structure between human and nonhuman bone. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Animal bone cross section (Figure 15.6)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The size of a bone can also help determine whether it belongs to a human. Adult human bones are larger than subadult or infant bones. However, another major difference between human adult bones and those of a young individual or infant human can be attributed to development and growth of the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1222\">epiphyses<\/a><\/strong> (ends of the bone). The epiphyses of human subadult bones are not fused to the shaft (Figure 15.7). Therefore, if a bone is small and it is suspected to belong to a human subadult or infant, the epiphyses would not be fused. Many small animal bones appear very similar in form compared to adult human bones, but they are much too small to belong to an adult human. Yet they can be eliminated as subadult or infant bones if the epiphyses are fused to the shaft.<\/p>\n<figure style=\"width: 288px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image13-3.png\" alt=\"X-ray image of child\u2019s ankle.\" width=\"288\" height=\"412\" \/><figcaption class=\"wp-caption-text\">Figure 15.7: An x-ray of a subadult\u2019s ankle with the epiphyses of the tibia and fibula visible. The gap between the shaft of the bone and the end of the bone (epiphysis) is the location of the growth plate. Therefore, the growth plate gap is what separates the shafts from the epiphyses in the image. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Tib_fib_growth_plates.jpg\">Tib fib growth plates<\/a> by <a href=\"https:\/\/en.wikipedia.org\/wiki\/User:Gilo1969\">Gilo1969<\/a> at <a href=\"https:\/\/en.wikipedia.org\/wiki\/\">English Wikipedia<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/3.0\/legalcode\">CC BY 3.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Is It Modern or Archaeological? <\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropologists work with modern cases that fall within the scope of law enforcement investigations. Accordingly, it is important to determine whether discovered human remains are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1224\">archaeological<\/a> <\/strong>or forensic in nature. Human remains that are historic are considered archeaological. The scientific study of human remains from archaeological sites is called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1226\">bioarchaeology<\/a><\/strong>.<\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Dig Deeper: Bioarchaeology<\/h2>\n<p class=\"import-Normal\">For readers who are interested in the sister subfield of bioarchaeology, which studies human remains and material culture from the past, please refer to chapter 8 of <em>Bioarchaeology: Interpreting Human Behavior from Skeletal Remains,<\/em> in <em>TRACES: An Open Invitation to Archaeology<\/em> (Blatt, Michael, and Bright forthcoming).<\/p>\n<\/div>\n<p>A forensic anthropologist should begin their analysis by reviewing the context in which the remains were discovered. This will help them understand a great deal about the remains, including determining whether they are archaeological or forensic in nature as well as considering legal and ethical issues associated with the collection, analysis, and storage of human remains (see \u201cEthics and Human Rights\u201d section of this chapter for more information).<\/p>\n<figure style=\"width: 403px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image10-3.png\" alt=\"Four teeth in a person\u2019s mouth. First molar with silver filling.\" width=\"403\" height=\"303\" \/><figcaption class=\"wp-caption-text\">Figure 15.8: A human tooth with a filling. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Filling.jpg#filehistory\">Filling<\/a> by Kauzio has been designated to the <a href=\"https:\/\/creativecommons.org\/share-your-work\/public-domain\/cc0\/\">public domain (CC0)<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The \u201ccontext\u201d refers to the relationship the remains have to the immediate area in which they were found. This includes the specific place where the remains were found, the soil or other organic matter immediately surrounding the remains, and any other objects or artifacts in close proximity to the body. For example, imagine that a set of remains has been located during a house renovation. The remains are discovered below the foundation. Do the remains belong to a murder victim? Or was the house built on top of an ancient burial ground? Observing information from the surroundings can help determine whether the remains are archaeological or modern. How long ago was the foundation of the house erected? Are there artifacts in close proximity to the body, such as clothing or stone tools? These are questions about the surroundings that will help determine the relative age of the remains.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Clues directly from the skeleton may also indicate whether the remains are archaeological or modern. For example, tooth fillings can suggest that the individual was alive recently (Figure 15.8). In fact, filling material has changed over the decades, so the specific type of material used to fix a cavity can be matched with specific time periods. Gold was used in dental work in the past, but more recently composite (a mixture of plastic and fine glass) fillings have become more common.<\/p>\n<h3><strong>How <\/strong><strong>Many Individuals Are Present?<\/strong><\/h3>\n<h4 class=\"import-Normal\"><em>What Is MNI?<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Another assessment that an anthropologist can perform is the calculation of the number of individuals in a mixed burial assemblage. Because not all burials consist of a single individual, it is important to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1268\">burial assemblage<\/a><\/strong> be able to estimate the number of individuals in a forensic context. Quantification of the number of individuals in a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1524\">burial assemblage<\/a><\/strong> can be done through the application of a number of methods, including the following: the Minimum Number of Individuals (MNI), the Most Likely Number of Individuals (MLNI), and the Lincoln Index (LI). The most commonly used method in biological anthropology, and the focus of this section, is determination of the MNI.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The MNI presents \u201cthe minimum estimate for the number of individuals that contributed to the sample\u201d (Adams and Konigsberg 2008, 243). Many methods of calculating MNI were originally developed within the field of zooarchaeology for use on calculating the number of individuals in faunal or animal assemblages (Adams and Konigsberg 2008, 241). What MNI calculations provide is a lowest possible count for the total number of individuals contributing to a skeletal assemblage. Traditional methods of calculating MNI include separating a skeletal assemblage into categories according to the individual bone and the side the bone comes from and then taking the highest count per category and assigning that as the minimum number (Figure 15.9).<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 664px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image28-3.png\" alt=\"Many bone portions laying on individual plastic bags on a table.\" width=\"664\" height=\"441\" \/><figcaption class=\"wp-caption-text\">Figure 15.9: Skeletal elements from a commingled faunal assemblage. Credit: Commingled animal remains from Eden-Farson Pre-Contact site in southwest Wyoming by Matt O\u2019Brien original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Why Calculate MNI?<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In a forensic context, the determination of MNI is most applicable in cases of mass graves, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1232\">commingled burials<\/a><\/strong>, and mass fatality incidents. The term <em>commingled<\/em> is applied to any burial assemblage in which individual skeletons are not separated into separate burials. As an example, the authors of this chapter have observed commingling of remains resulting from mass fatality wildfire events. Commingled remains may also be encountered in events such as a plane or vehicle crash. It is important to remember that in any forensic context, MNI should be referenced and an MNI of one should be substantiated by the fact that there was no repetition of elements associated with the case.<\/p>\n<h3 class=\"import-Normal\"><strong>Constructing the Biological Profile<\/strong><\/h3>\n<h4 class=\"import-Normal\"><em>Who Is It?<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">\u201cWho is it?\u201d is one of the first questions that law enforcement officers ask when they are faced with a set of skeletal remains. To answer this question, forensic anthropologists construct a biological profile (White and Folkens 2005, 405). A <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1228\">biological profile<\/a> <\/strong>is an individual\u2019s identifying characteristics, or biological information, which include the following: biological sex, age at death, stature, population affinity, skeletal variation, and evidence of trauma and pathology.<\/p>\n<h4 class=\"import-Normal\"><em>Assessing Biological Sex <\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Assessment of biological sex is often one of the first things considered when establishing a biological profile because several other parts, such as age and stature estimations, rely on an assessment of biological sex to make the calculations more accurate.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Assessment of biological sex focuses on differences in both morphological (form or structure) and metric (measured) traits in individuals. When assessing morphological traits, the skull and the pelvis are the most commonly referenced areas of the skeleton. These differences are related to sexual dimorphism usually varying in the amount of robusticity seen between males and females. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1230\">Robusticity<\/a> <\/strong>deals with strength and size; it is frequently used as a term to describe a large size or thickness. In general, males will show a greater degree of robusticity than females. For example, the length and width of the mastoid process, a bony projection located behind the opening for the ear, is typically larger in males. The mastoid process is an attachment point for muscles of the neck, and this bony projection tends to be wider and longer in males. In general, cranial features tend to be more robust in males (Figure 15.10).<\/p>\n<figure style=\"width: 601px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image26-3.png\" alt=\"Front and side images of a male (left) and female (right) cranium.\" width=\"601\" height=\"632\" \/><figcaption class=\"wp-caption-text\">Figure 15.10: Anterior and lateral view of a male and female cranium. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Anterior and lateral view of a male and female cranium (Figure 15.10)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropo logy<\/a> by Ashley Kendell is a collective work under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Includes <a href=\"https:\/\/boneclones.com\/product\/modern-human-asian-female-skull-BC-149\/category\/all-human-skulls\/human-anatomy\">Human Female Asian Skull<\/a> and <a href=\"https:\/\/boneclones.com\/product\/human-asian-male-skull-BC-016\/category\/all-human-skulls\/human-anatomy\">Human Male Asian Skull<\/a> by <a href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a>, used by permission.]<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">When considering the pelvis, the features associated with the ability to give birth help distinguish females from males. During puberty, estrogen causes a widening of the female pelvis to allow for the passage of a baby. Several studies have identified specific features or bony landmarks associated with the widening of the hips, and this section will discuss one such method. The Phenice Method (Phenice 1969) is traditionally the most common reference used to assess morphological characteristics associated with sex. The Phenice Method specifically looks at the presence or absence of (1) a ventral arc, (2) the presence or absence of a subpubic concavity, and (3) the width of the medial aspect of the ischiopubic ramus (Figure 15.11). When present, the ventral arc, a ridge of bone located on the ventral surface of the pubic bone, is indicative of female remains. Likewise the presence of a subpubic concavity and a narrow medial aspect of the ischiopubic ramus is associated with a female sex estimation. Assessments of these features, as well as those of the skull (when both the pelvis and skull are present), are combined for an overall estimation of sex.<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 1603px\" class=\"wp-caption alignnone\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image29-3.png\" alt=\"Male and female os coxae (anterior portions).\" width=\"1603\" height=\"582\" \/><figcaption class=\"wp-caption-text\">Figure 15.11: Features associated with the Phenice Method. Images derived from CSU-HIL donated skeletal collection. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Features associated with the Phenice Method (Figure 15.11)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Colleen Milligan is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Metric analyses are also used in the estimation of sex. Measurements taken from every region of the body can contribute to estimating sex through statistical approaches that assign a predictive value of sex. These approaches can include multiple measurements from several skeletal elements in what is called multivariate (multiple variables) statistics. Other approaches consider a single measurement, such as the diameter of the head of the femur, of a specific element in a univariate (single variable) analysis (Berg 2017, 152\u2013156).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">It is important to note that, although forensic anthropologists usually begin assessment of biological profile with biological sex, there is one major instance in which this is not appropriate. The case of two individuals found in California, on July 8, 1979, is one example that demonstrates the effect age has on the estimation of sex. The identities of the two individuals were unknown; therefore, law enforcement sent them to a lab for identification. A skeletal analysis determined that the remains represented one adolescent male and one adolescent female, both younger than 18 years of age. This information did not match with any known missing children at the time.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In 2015, the cold case was reanalyzed, and DNA samples were extracted. The results indicated that the remains were actually those of two girls who went missing in 1978. The girls were 15 years old and 14 years old at the time of death. It is clear that the 1979 results were incorrect, but this mistake also provides the opportunity to discuss the limitations of assessing sex from a subadult skeleton.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Assessing sex from the human skeleton is based on biological and genetic traits associated with females and males. These traits are linked to differences in sexual dimorphism and reproductive characteristics between females and males. The link to reproductive characteristics means that most indicators of biological sex do not fully manifest in prepubescent individuals, making estimations of sex unreliable in younger individuals (SWGANTH 2010b). This was the case in the example of the 14-year-old girl. When examined in 1979, her remains were misidentified as male because she had not yet fully developed female pelvic traits.<\/p>\n<h4 class=\"import-Normal\"><em>Sex vs. Gender<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Biological sex is a different concept than <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1234\">gender<\/a><\/strong>. While biological anthropologists can estimate sex from the skeleton, estimating an individual\u2019s gender would require a greater context because gender is defined culturally rather than biologically. Take, for example, an individual who identifies as transgender. This individual has a gender identity that is different from their biological sex. The gender identity of any individual depends on factors related to self-identification, situation or context, and cultural factors. <span style=\"background-color: #00ffff\">While in the U.S<\/span>. we have historically thought of sex and gender as binary concepts (male or female), many cultures throughout the world recognize several possible gender identities. In this sense, gender is seen as a continuous or fluid variable rather than a fixed one.<\/p>\n<p class=\"import-Normal\">Historically, forensic anthropologists have used a binary construct to categorize human skeletal remains as either male or female (with the accompanying categories of probable male, probable female, and indeterminate). In the case of transgender and gender nonconforming individuals, the binary approach to sex assessment may delay or hinder identification efforts (Buchanan 2014; Schall, Rogers, and Deschamps-Braly 2020; Tallman, Kincer, and Plemons 2021). As such, many forensic anthropologists have begun to address the inherent problems associated with a binary approach to sex identification and to explore ways of assessing social identity and self-identified gender using skeletal remains and forensic context.<\/p>\n<p class=\"import-Normal\">For the duration of this section, the term <em>transgender<\/em> refers to individuals whose gender identity differs from the sex assigned at birth (Schall, Rogers, and Deschamps-Braly 2020:2). Transgender individuals transition from one gender binary to another, such as male-to-female (MTF) or female-to-male (FTM). While many of the gender-affirming procedures available to trans and gender-nonconforming individuals are focused on soft tissue modifications (e.g., breast augmentation, genital reconstruction, hormone therapies, etc.), there are a number of gender-affirmation surgeries that do leave a permanent record on the skeleton. Generally speaking, FTM transgender people are reported to undergo fewer surgical procedures than do MTF transgender people (Buchanan 2014). The discussion below focuses on Facial Feminization Surgery (FFS), which leaves a permanent record on the human skeleton that may be used to help make an identification.<\/p>\n<p class=\"import-Normal\">FFS refers to a combination of procedures focused on sexually dimorphic features of the face, with the intent of transforming typically male facial features into more feminine forms. Facial Feminization Surgery procedures were developed by Dr. Douglas Ousterhout, a San Francisco based cranio-maxillofacial surgeon, in the mid-1980s (Schall, Rogers, and Deschamps-Braly 2020:2). FFS can include one or a combination of the following: hairline lowering, forehead reduction and contouring, brow lift, reduction rhinoplasty, cheek enhancement, lift lift, lip filling, chin contouring, jaw contouring, and\/or tracheal shave (Buchanan 2014; Schall, Rogers, and Deschamps-Braly 2020:2). Of the procedures outlined previously, four are known to directly affect the facial skeleton: forehead contouring, rhinoplasty, chin contouring, and jaw contouring (Buchanan 2014; Schall, Rogers, and Deschamps-Braly 2020:2).<\/p>\n<p class=\"import-Normal\">Because FFS procedures have been widely documented in the medical (and more recently the forensic anthropological) literature, there are a number of indicators that a forensic anthropologist can use to make more informed evaluations of gender, including evidence of bone remodeling in sexually dimorphic regions of the skull (e.g., forehead, chin, jawline), as well as the presence of plates, pins, or other surgical hardware that may be evidence of FFS (Buchanan 2014; Schall, Rogers, and Deschamps-Braly 2020; Tallman, Kincer, and Plemons 2021). Additionally, some forensic anthropologists suggest cautiously integrating contextual information from the scene, such as personal effects, material evidence, and recovery scene information, into their evaluation of an individual\u2019s social identity (Beatrice and Soler 2016; Birkby, Fenton, and Anderson 2008; Soler and Beatrice 2018; Soler et al. 2019; Tallman, Kincer, and Plemons 2021; Winburn, Schoff, and Warren 2016). The ultimate goal of many skeletal analyses is to make a positive identification on a set of unidentified remains.<\/p>\n<h4 class=\"import-Normal\"><em>Assessment <\/em><em>of Population Affinity<\/em><\/h4>\n<p>In an effort to combat the erroneous assumptions tied to the race concept, forensic anthropologists have attempted to reframe this component of the biological profile. The term <em>race<\/em> is no longer used in casework and teaching. Historically, the word <em>ancestry<\/em> is and was deemed a more appropriate way to describe an individual\u2019s phenotype. However, in more recent years, forensic anthropologists have begun using the term <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1236\">population affinity<\/a><\/strong><em>, <\/em>recognizing that we are basing our analysis on the similarities we see based on the reference samples we have available (Winburn and Algee-Hewitt 2021). An important note here is that it is possible to hinder identifications and harm individuals when tools like estimations of population affinity are misapplied, misinterpreted, or misused. For this reason, the field of forensic anthropology has ongoing conversations about the appropriateness of this analysis in the biological profile (Bethard and DiGangi 2020; Stull et al. 2021).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">We use the term <em>population affinity<\/em> to refer to the variation seen among modern populations\u2014variation that is both genetic and environmentally driven. The word <em>affinity<\/em> refers to similarities or relationships between individuals. As forensic anthropologists, we compare an unknown individual to multiple reference groups and look for the degree of similarity in observable traits with those groups. As noted previously, population affinity can aid law enforcement in their identification of missing persons or unknown skeletal remains.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Within the field of anthropology, the estimation of population affinity has a contentious history, and early attempts at classification were largely based on the erroneous assumption that an individual\u2019s <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1238\">phenotype <\/a><\/strong> (outward appearance) was correlated with their innate intelligence and abilities (see Chapter 13 for a more in-depth discussion of the history of the race concept). The use of the term <em>race<\/em> is deeply embedded in the social context of the United States. In any other organism\/living thing, groups divided according to the biological race concept would be defined as a separate subspecies. The major issue with applying the biological race concept to humans is that there are not enough differences between any two populations to separate on a genetic basis. In other words, <em>biological races do not exist in human populations. <\/em>However, the concept of race has been perpetuated and upheld by sociocultural constructs of race.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The conundrum for forensic anthropologists is the fact that while races do not exist on a biological level, we still socially recognize and categorize individuals based on their phenotype. Clearly, our phenotype is an important factor in not only how we are viewed by others but also how we identify ourselves. It is also a commonly reported variable. Often labeled as \u201crace,\u201d we are asked to report how we self-identify on school applications, government identification, surveys, census reports, and so forth. It follows then that when a person is reported missing, the information commonly collected by law enforcement and sometimes entered into a missing person\u2019s database includes their age, biological sex, stature, and \u201crace.\u201d Therefore, the more information a forensic anthropologist can provide regarding the individual\u2019s physical characteristics, the more he or she can help to narrow the search.<\/p>\n<p class=\"import-Normal\">As an exercise, create a list of all of the women you know who are between the ages of 18 and 24 and approximately 5\u2019 4\u201d to 5\u2019 9\u201d tall. You probably have several dozen people on the list. Now, consider how many females you know who are between the ages of 18 and 24, are approximately 5\u2019 4\u201d to 5\u2019 9\u201d tall, and are Vietnamese. Your list is going to be significantly shorter. That\u2019s how missing persons searches go as well. The more information you can provide regarding a decedent\u2019s phenotype, the fewer possible matches law enforcement are left to investigate. This is why population affinity has historically been included as a part of the biological profile.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Traditionally, population affinity was accomplished through a visual inspection of morphological variants of the skull (morphoscopics). These methods focused on elements of the facial skeleton, including the nose, eyes, and cheek bones. However, in an effort to reduce subjectivity, nonmetric cranial traits are now assessed within a statistical framework to help anthropologists better interpret their distribution among living populations (Hefner and Linde 2018). Based on the observable traits, a macromorphoscopic analysis will allow the practitioner to create a statistical prediction of geographic origin. In essence, forensic anthropologists are using human variation in the estimation of geographic origin, by referencing documented frequencies of nonmetric skeletal indicators or macromorphoscopic traits.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Population affinity is also assessed through metric analyses. The computer program Fordisc is an anthropological tool used to estimate different components of the biological profile, including ancestry, sex, and stature. When using Fordisc, skeletal measurements are input into the computer software, and the program employs multivariate statistical classification methods, including discriminant function analysis, to generate a statistical prediction for the geographic origin of unknown remains based on the comparison of the unknown to the reference samples in the software program. Fordisc also calculates the likelihood of the prediction being correct, as well as how typical the metric data is for the assigned group.<\/p>\n<h4 class=\"import-Normal\"><em>Estimating Age-at-Death<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Estimating age-at-death from the skeleton relies on the measurement of two basic physiological processes: (1) growth and development and (2) degeneration (or aging). From fetal development on, our bones and teeth grow and change at a predictable rate. This provides for relatively accurate age estimates. After our bones and teeth cease to grow and develop, they begin to undergo structural changes, or degeneration, associated with aging. This does not happen at such predictable rates and, therefore, results in less accurate or larger age-range estimations.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">During growth and development stages, two primary methods used for estimations of age of subadults (those under the age of 18) are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1240\">epiphyseal union<\/a><\/strong> and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1244\">dental development.<\/a><\/strong> Epiphyseal union<strong> (<\/strong>or <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1242\">epiphyseal fusion<\/a><\/strong>) refers to the appearance and closure of the epiphyseal plates between the primary centers of growth in a bone and the subsequent centers of growth (see Figure 15.7). Prior to complete union, the cartilaginous area between the primary and secondary centers of growth is also referred to as the growth plates (Schaefer, Black, and Scheuer 2009). Different areas of the skeleton have documented differences in the appearance and closure of epiphyses, making this a reliable method for aging subadult remains (SWGANTH 2013).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">As an example of its utility in the identification process, epiphyseal development was used to identify two subadult victims of a fatal fire in Flint, Michigan, in February 2010. The remains represented two young girls, ages three and four. Due to the intensity of the fire, the subadult victims were differentiated from each other through the appearance of the patella, the kneecap. The patella is a bone that develops within the tendon of the quadriceps muscle at the knee joint. The patella begins to form around three to four years of age (Cunningham, Scheuer, and Black 2016, 407\u2013409). In the example above, radiographs of the knees showed the presence of a patella in the four-year-old girl and the absence of a clearly discernible patella in the three-year-old.<\/p>\n<figure style=\"width: 358px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image18-2.png\" alt=\"Cranial cast of child with exposed maxilla and mandible to see developing dentition.\" width=\"358\" height=\"358\" \/><figcaption class=\"wp-caption-text\">Figure 15.12: Dental development in a subadult. Credit: <a href=\"https:\/\/boneclones.com\/product\/5-year-old-human-child-skull-with-mixed-dentition-exposed-BC-189\">5-year-old Human Child Skull with Mixed Dentition Exposed<\/a> by <a href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a> is used by permission and available here is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Dental development begins during fetal stages of growth and continues until the complete formation and eruption of the adult third molars (if present). The first set of teeth to appear are called deciduous or baby teeth. Individuals develop a total of 20 deciduous teeth, including incisors, canines, and molars. These are generally replaced by adult dentition as an individual grows (Figure 15.12). A total of 32 teeth are represented in the adult dental arcade, including incisors, canines, premolars, and molars. When dental development is used for age estimations, researchers use both tooth-formation patterns and eruption schedules as determining evidence. For example, the crown of the tooth forms first followed by the formation of the tooth root. During development, an individual can exhibit a partially formed crown or a complete crown with a partially formed root. The teeth generally begin the eruption process once the crown of the tooth is complete. The developmental stages of dentition are one of the most reliable and consistent aging methods for subadults (Langley, Gooding, and Tersigni-Tarrant 2017, 176\u2013177).<\/p>\n<figure style=\"width: 403px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image22-3.png\" alt=\"Surfaces of three pubic symphyses: billowy (A) to more flat (B) to rough (C).\" width=\"403\" height=\"224\" \/><figcaption class=\"wp-caption-text\">Figure 15.13: Examples of degenerative changes to the pubic symphysis: (A) young adult; (B) middle adult; (C) old adult. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Example of the progression of degenerative changes to the pubic symphysis (Figure 15.14)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropo logy<\/a> by Ashley Kendell is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Original photos by Dr. Julie Fleischman used by permission. Pubic symphyses are curated in the Hartnett-Fulginiti donated skeletal collection. Donation and research consent was provided by next of kin.]<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Degenerative changes in the skeleton typically begin after 18 years of age, with more prominent changes developing after an individual reaches middle adulthood (commonly defined as after 35 years of age in osteology). These changes are most easily seen around joint surfaces of the pelvis, the cranial vault, and the ribs. In this chapter, we focus on the pubic symphysis surfaces of the pelvis and the sternal ends of the ribs, which show metamorphic changes from young adulthood to older adulthood. The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1262\">pubic symphysis<\/a> <\/strong>is a joint that unites the left and right halves of the pelvis. The surface of the pubic symphysis changes during adulthood, beginning as a surface with pronounced ridges (called billowing) and flattening with a more distinct rim to the pubic symphysis as an individual ages. As with all metamorphic age changes, older adults tend to develop lipping around the joint surfaces as well as a breakdown of the joint surfaces. The most commonly used method for aging adult skeletons from the pubic symphysis is the Suchey-Brooks method (Brooks and Suchey 1990; Katz and Suchey 1986). This method divides the changes seen with the pubic symphysis into six phases based on macroscopic age-related changes to the surface. Figure 15.13 provides a visual of the degenerative changes that typically occur on the pubic symphysis.<\/p>\n<figure style=\"width: 403px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image12-3.png\" alt=\"Three sternal rib ends demonstrating progressive changes that occur with age.\" width=\"403\" height=\"220\" \/><figcaption class=\"wp-caption-text\">Figure 15.14: Examples of degenerative changes to the sternal rib end: (A) young adult; (B) middle adult; (C) old adult. Images derived from CSU, Chico HIL donated skeletal collection. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Examples of degenerative changes to the sternal rib end (Figure 15.15)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The sternal end of the ribs, the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1246\">anterior <\/a><\/strong> end of the rib that connects via cartilage to the sternum, is also used in age estimations of adults. This method, first developed by M. Y. \u0130\u015fcan and colleagues, considers both the change in shape of the sternal end as well as the quality of the bone (\u0130\u015fcan, Loth, and Wright 1984; \u0130\u015fcan, Loth, and Wright 1985). The sternal end first develops a billowing appearance in young adulthood. The bone typically develops a wider and deeper cupped end as an individual ages. Older adults tend to exhibit bony extensions of the sternal end rim as attaching cartilage ossifies. Figure 15.14 provides a visual of the degenerative changes that typically occur in sternal rib ends.<\/p>\n<h4 class=\"import-Normal\"><em>Estimating Stature<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Stature, or height, is one of the most prominently recorded components of the biological profile. Our height is recorded from infancy through adulthood. Doctor\u2019s appointments, driver's license applications, and sports rosters all typically involve a measure of stature for an individual. As such, it is also a component of the biological profile nearly every individual will have on record. Bioarchaeologists and forensic anthropologists use stature estimation methods to provide a range within which an individual\u2019s biological height would fall. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1248\">Biological height<\/a> <\/strong>is a person\u2019s true anatomical height. However, the range created through these estimations is often compared to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1264\">reported stature<\/a><\/strong>, which is typically self-reported and based on an approximation of an individual\u2019s true height (Ousley 1995).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In June 2015, two men were shot and killed in Granite Bay, California, in a double homicide. Investigators were able to locate surveillance camera footage from a gas station where the two victims were spotted in a car with another individual believed to be the perpetrator in the case. The suspect, sitting behind the victims in the car, hung his right arm out of the window as the car drove away. The search for the perpetrator was eventually narrowed down to two suspects. One suspect was 5\u2019 8\u201d while the other suspect was 6\u2019 4\u201d, representing almost a foot difference in height reported stature between the two. Forensic anthropologists were given the dimensions of the car (for proportionality of the arm) and were asked to calculate the stature of the suspect in the car from measurements of the suspect\u2019s forearm hanging from the window. Approximate lengths of the bones of the forearm were established from the video footage and used to create a predicted stature range. Stature estimations from skeletal remains typically look at the correlation between the measurements of any individual bone and the overall measurement of body height. In the case above, the length of the right forearm pointed to the taller of the two suspects who was subsequently arrested for the homicide.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Certain bones, such as the long bones of the leg, contribute more to our overall height than others and can be used with mathematical equations known as regression equations. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1252\">Regression methods <\/a> <\/strong>examine the relationship between variables such as height and bone length and use the correlation between the variables to create a prediction interval (or range) for estimated stature. This method for calculating stature is the most commonly used method (SWGANTH 2012). Figure 15.15 shows the measurement of the bicondylar length of the femur for stature estimations.<\/p>\n<figure style=\"width: 584px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image2-3.png\" alt=\"A femur is measured using a wooden osteometric board.\" width=\"584\" height=\"389\" \/><figcaption class=\"wp-caption-text\">Figure 15.15: Image of measurement of the bicondylar length of the femur, often used in the estimation of living stature. Image derived from CSU, Chico HIL donated skeletal collection. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Measurement of the bicondylar length of the femur<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Identification Using Individualizing Characteristics<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">One of the most frequently requested analyses within the forensic anthropology laboratory is assistance with the identification of unidentified remains. While all components of a biological profile, as discussed above, can assist law enforcement officers and medical examiners to narrow down the list of potential identifications, a biological profile will not lead to a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1254\">positive identification<\/a><\/strong>. The term <em>positive identification<\/em> refers to a scientifically validated method of identifying previously unidentified remains. Presumptive identifications, however, are not scientifically validated; rather, they are based on circumstances or scene context. For example, if a decedent is found in a locked home with no evidence of forced entry but the body is no longer visually identifiable, it may be presumed that the remains belong to the homeowner. Hence, a presumptive identification.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The medicolegal system ultimately requires that a positive identification be made in such circumstances, and a presumptive identification is often a good way to narrow down the pool of possibilities. Biological profile information also assists with making a presumptive identification based on an individual\u2019s phenotype in life (e.g., what they looked like). As an example, a forensic anthropologist may establish the following components of a biological profile: white male, between the ages of 35 and 50, approximately 5\u2019 7\u201d to 5\u2019 11.\u201d While this seems like a rather specific description of an individual, you can imagine that this description fits dozens, if not hundreds, of people in an urban area. Therefore, law enforcement can use the biological profile information to narrow their pool of possible identifications to include only white males who fit the age and height outlined above. Once a possible match is found, the decedent can be identified using a method of positive identification.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Positive identifications are based on what we refer to as individualizing traits or characteristics, which are traits that are unique at the individual level. For example, brown hair is not an individualizing trait as brown is the most common hair color in the U.S. But, a specific pattern of dental restorations or surgical implants can be individualizing, because it is unlikely that you will have an exact match on either of these traits when comparing two individuals.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">A number of positive methods are available to forensic anthropologists, and for the remainder of this section we will discuss the following methods: comparative medical and dental radiography and identification of surgical implants.<\/p>\n<figure style=\"width: 165px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image17-3.png\" alt=\"Radiograph of skull with frontal sinuses visible.\" width=\"165\" height=\"182\" \/><figcaption class=\"wp-caption-text\">Figure 15.16: Example of the unique shape of the frontal sinus. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Frontal_bone_sinuses.jpg\">Frontal bone sinuses<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Alex_Khimich\">Alex Khimich<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Comparative medical and dental radiography is used to find consistency of traits when comparing antemortem records (medical and dental records taken during life) with images taken postmortem (after death). Comparative medical radiography focuses primarily on features associated with the skeletal system, including trabecular pattern (internal structure of bone that is honeycomb in appearance), bone shape or cortical density (compact outer layer of bone), and evidence of past trauma, skeletal pathology, or skeletal anomalies. Other individualizing traits include the shape of various bones or their features, such as the frontal sinuses (Figure 15.16).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Comparative dental radiography focuses on the number, shape, location, and orientation of dentition and dental restorations in antemortem and postmortem images. While there is not a minimum number of matching traits that need to be identified for an identification to be made, the antemortem and postmortem records should have enough skeletal or dental consistencies to conclude that the records did in fact come from the same individual (SWGANTH 2010a). Consideration should also be given to population-level frequencies of specific skeletal and dental traits. If a trait is particularly common within a given population, it may not be a good trait to utilize for positive identification.<\/p>\n<figure style=\"width: 354px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image16-3.png\" alt=\"A scapula and humerus with a metal shoulder replacement.\" width=\"354\" height=\"231\" \/><figcaption class=\"wp-caption-text\">Figure 15.17: Image of joint replacement in the right shoulder. Credit: <a href=\"https:\/\/naturalhistory.si.edu\/education\/teaching-resources\/written-bone\/skeleton-keys\/todays-bones\">Shoulder replacement<\/a> by <a href=\"https:\/\/www.si.edu\/\">Smithsonian<\/a> [exhibit: Written in Bone, Today\u2019s Bones] <a href=\"https:\/\/www.si.edu\/termsofuse\">is used for educational and non-commercial purposes as outlined by the Smithsonian.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Surgical implants or devices can also be used for identification purposes (Figure 15.17). These implements are sometimes recovered with human remains. One of the ways forensic anthropologists can use surgical implants to assist in decedent identification is by providing a thorough analysis of the implant and noting any identifying information such as serial numbers, manufacturer symbols, and so forth. This information can then sometimes be tracked directly to the manufacturer or the place of surgical intervention, which may be used to identify unknown remains (SWGANTH 2010a).<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Trans Doe Task Force<\/h2>\n<p class=\"import-Normal\">The Trans Doe Task Force (TDTF) is a Trans-led nonprofit organization that investigates cases involving LGBTQ+ missing and murdered persons. The organization specifically focuses on transgender and gender-variant cases, providing connections between law enforcement agencies, medical examiner offices, forensic anthropologists, and forensic genetic genealogists to increase the chances of identification. Additionally, the TDTF curates a data repository of missing, murdered, and unclaimed LGBTQ+ individuals, and they continuously try innovative approaches to identify these individuals, whose lived gender identity may not match their biological sex.<\/p>\n<p class=\"import-Normal\">For more information visit <a href=\"https:\/\/transdoetaskforce.org\/\">transdoetaskforce.org<\/a><\/p>\n<\/div>\n<h3 class=\"import-Normal\"><strong>Trauma Analysis<\/strong><\/h3>\n<h4 class=\"import-Normal\"><em>Types of Trauma<\/em><strong><br \/>\n<\/strong><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Within the field of anthropology, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1256\">trauma<\/a> <\/strong>is defined as an injury to living tissue caused by an extrinsic force or mechanism (Lovell 1997:139). Forensic anthropologists can assist a forensic pathologist by providing an interpretation of the course of events that led to skeletal trauma. Typically, traumatic injury to bone is classified into one of four categories, defined by the trauma mechanism. A trauma mechanism refers to the force that produced the skeletal modification and can be classified as (1) sharp force, (2) blunt force, (3) projectile, or (4) thermal (burning). Each type of trauma, and the characteristic pattern(s) associated with that particular categorization, will be discussed below.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">First, let\u2019s consider s<em>harp-force trauma<\/em>, which is caused by a tool that is edged, pointed, or beveled\u2014for example, a knife, saw, or machete (SWGANTH 2011). The patterns of injury resulting from sharp-force trauma include linear incisions created by a sharp, straight edge; punctures; and chop marks (Figure 15.18; SWGANTH 2011). When observed under a microscope, an anthropologist can often determine what kind of tool created the bone trauma. For example, a power saw cut will be discernible from a manual saw cut.<\/p>\n<figure style=\"width: 602px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image21-1.png\" alt=\"Anterior image of a skull with multiple traumatic injuries to forehead.\" width=\"602\" height=\"457\" \/><figcaption class=\"wp-caption-text\">Figure 15.18: Example of sharp-force trauma (sword wound) to the frontal bone. The skull appears sliced with thin lines in two places across the top of the skull. Credit: <a href=\"https:\/\/openverse.org\/image\/909d1b77-ad5f-4cda-be44-6d9b5fbf14b9\/\">Female skull injured by a medieval sword<\/a> by <a href=\"https:\/\/sketchfab.com\/provinciaal_depot_noordholland\">Provinciaal depot voor archeologie Noord-Holland<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY 4.0 License<\/a>. The original image is a 3D model that can be manipulated on the <a href=\"https:\/\/wordpress.org\/openverse\/image\/909d1b77-ad5f-4cda-be44-6d9b5fbf14b9\/\">openverse website<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Second, <em>blunt-force trauma<\/em> is defined as \u201ca relatively low-velocity impact over a relatively large surface area\u201d (Galloway 1999, 5). Blunt-force injuries can result from impacts from clubs, sticks, fists, and so forth. Blunt-force impacts typically leave an injury at the point of impact but can also lead to bending and deformation in other regions of the bone. Depressions, fractures, and deformation at and around the site of impact are all characteristics of blunt-force trauma (Figure 15.19). As with sharp-force trauma, an anthropologist attempts to interpret blunt-force injuries, providing information pertaining to the type of tool used, the direction of impact, the sequence of impacts, if more than one, and the amount of force applied.<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 578px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image30.png\" alt=\"Cranium with two blunt force impacts from a hammer.\" width=\"578\" height=\"803\" \/><figcaption class=\"wp-caption-text\">Figure 15.19: Example of multiple blunt force impacts to the left parietal and frontal bones. There is one hole in the skull with fractured bone around the edges. There are also multiple spots across the back of the skull with depressions of various sizes. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Skull_hammer_trauma.jpg\">Skull hammer trauma<\/a> by <a href=\"https:\/\/www.nih.gov\/\">the National Institutes of Health<\/a>, Health &amp; Human Services, is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>. [Exhibit: Visible Proofs: Forensic Views of the Body, U.S. National Library of Medicine, 19th Century Collection, National Museum of Health and Medicine, Armed Forces Institute of Pathology, Washington, D.C.]<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Third, <em>projectile trauma<\/em> refers to high-velocity trauma, typically affecting a small surface area (Galloway 1999, 6). Projectile trauma results from fast-moving objects such as bullets or shrapnel. It is typically characterized by penetrating defects or embedded materials (Figure 15.20). When interpreting injuries resulting from projectile trauma, an anthropologist can often offer information pertaining to the type of weapon used (e.g., rifle vs. handgun), relative size of the bullet (but not the caliber of the bullet), the direction the projectile was traveling, and the sequence of injuries if there are multiple present.<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 462px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image5-3.png\" alt=\"Anterior and posterior views of a skull with a gunshot wound.\" width=\"462\" height=\"291\" \/><figcaption class=\"wp-caption-text\">Figure 15.20: Example of projectile trauma with an entrance wound to the frontal bone and exit wound visible on the occipital. A small circular hole is visible in the front of the skull with cracks radiating out from the point of impact. There is a larger hole visible in the back of the skull that is irregular yet circular in shape. Credit: <a href=\"https:\/\/naturalhistory.si.edu\/education\/teaching-resources\/written-bone\/skeleton-keys\/how-bone-biographies-get-written\">Trauma: Gunshot Wounds<\/a> by <a href=\"https:\/\/www.si.edu\/\">Smithsonian<\/a> [exhibit: Written in Bone, How Bone Biographies Get Written] <a href=\"https:\/\/www.si.edu\/termsofuse\">is used for educational and non-commercial purposes as outlined by the Smithsonian.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Finally, <em>thermal trauma<\/em> is a bone alteration that results from bone exposure to extreme heat. Thermal trauma can result in cases of house or car fires, intentional disposal of a body in cases of homicidal violence, plane crashes, and so on. Thermal trauma is most often characterized by color changes to bone, ranging from yellow to black (charred) or white (calcined). Other bone alterations characteristic of thermal trauma include delamination (flaking or layering due to bone failure), shrinkage, fractures, and heat-specific burn patterning. When interpreting injuries resulting from thermal damage, an anthropologist can differentiate between thermal fractures and fractures that occurred before heat exposure, thereby contributing to the interpretation of burn patterning (e.g., was the individual bound or in a flexed position prior to the fire?).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">While there are characteristic patterns associated with the four categories of bone trauma, it is also important to note that these bone alterations do not always occur independently of different trauma types. An individual\u2019s skeleton may present with multiple different types of trauma, such as a projectile wound and thermal trauma. Therefore, it is important that the anthropologist recognize the different types of trauma and interpret them appropriately.<\/p>\n<h3 class=\"import-Normal\"><strong>Timing of Injury<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Another important component of any anthropological trauma analysis is the determination of the timing of injury (e.g., when did the injury occur). Timing of injury is traditionally split into one of three categories: <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1260\">antemortem<\/a> <\/strong>(before death), <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1258\">perimortem<\/a> <\/strong>(at or around the time of death), and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1266\">postmortem <\/a><\/strong>(after death). This classification system differs slightly from the classification system used by the pathologist because it specifically references the qualities of bone tissue and bone response to external forces. Therefore, the perimortem interval (at or around the time of death) means that the bone is still fresh and has what is referred to as a green bone response, which can extend past death by several weeks or even months. For example, in cold or freezing temperatures a body can be preserved for extended periods of time, increasing the perimortem interval, while in desert climates decomposition is accelerated, thereby significantly decreasing the postmortem interval (Galloway 1999, 12). Antemortem injuries (occurring well before death and not related to the death incident) are typically characterized by some level of healing, in the form of a fracture callus or unification of fracture margins. Finally, postmortem injuries (occurring after death, while bone is no longer fresh) are characterized by jagged fracture margins, resulting from a loss of moisture content during the decomposition process (Galloway 1999, 16). In general, all bone traumas should be classified according to the timing of injury, if possible. This information will help the medical examiner or pathologist better understand the circumstances surrounding the decedent\u2019s death, as well as events occurring during life and after the final disposition of the body.<\/p>\n<h3 class=\"import-Normal\"><strong>The Role of the Forensic Anthropologist in Trauma Analysis<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Within the medicolegal system, forensic anthropologists are often called upon by the medical examiner, forensic pathologist, or coroner to assist with an interpretation of trauma. The forensic anthropologist\u2019s main focus in any trauma analysis is the underlying skeletal system\u2014as well as, sometimes, cartilage. Analysis and interpretation of soft tissue injuries fall within the purview of the medical examiner or pathologist. It is also important to note that the main role of the forensic anthropologist is to provide information pertaining to skeletal injury to assist the medical examiner\/pathologist in their final interpretation of injury. Forensic anthropologists do not hypothesize as to the cause of death of an individual. Instead, a forensic anthropologist\u2019s report should include a description of the injury (e.g., trauma mechanism, number of injuries, location, timing of injury); documentation of the injury, which may be utilized in court testimony (e.g., photographs, radiographs, measurements); and, if applicable, a statement as to the condition of the body and state of decomposition, which may be useful for understanding the depositional context (e.g., how long has the body been exposed to the elements; was it moved or in its original location; are any of the alterations to bone due to environmental or faunal exposure instead of intentional human modification).<\/p>\n<h2 class=\"import-Normal\">Taphonomy<\/h2>\n<h2 class=\"import-Normal\"><strong>What Happened to the Remains After Death?<\/strong><\/h2>\n<p class=\"import-Normal\">The majority of the skeletal analysis process revolves around the identity of the deceased individual. However, there is one last, very important question that forensic anthropologists should ask: What happened to the remains after death? Generally speaking, processes that alter the bone after death are referred to as taphonomic changes (refer to Chapter 7 for a discussion regarding taphonomy and the fossil record).<\/p>\n<p class=\"import-Normal\">The term <em>taphonomy<\/em> was originally used to refer to the processes through which organic remains mineralize, also known as fossilization. Within the context of biological anthropology, the term <em>taphonomy<\/em> is better defined as the study of what happens to human remains after death (Komar and Buikstra 2008). Initial factors affecting a body after death include processes such as decomposition and scavenging by animals. However, taphonomic processes encompass much more than the initial period after death. For example, plant root growth can leach minerals from bone, leaving a distinctive mark. Sunlight can bleach human remains, leaving exposed areas whiter than those that remained buried. Water can wear the surface of the bone until it becomes smooth.<\/p>\n<p class=\"import-Normal\">Some taphonomic processes can help a forensic anthropologist estimate the relative amount of time that human remains have been exposed to the elements. For example, root growth through a bone would certainly indicate a body was buried for more than a few days. Forensic anthropologists must be very careful when attempting to estimate time since death based on taphonomic processes because environmental conditions can greatly influence the rate at which taphonomic processes progress. For example, in cold environments, tissue may decay slower than in warm, moist environments.<\/p>\n<p class=\"import-Normal\">Forensic anthropologists must contend with taphonomic processes that affect the preservation of bones. For example, high acidity in the soil can break down human bone to the point of crumbling. In addition, when noting trauma, they must be very careful not to confuse postmortem (after death) bone damage with trauma.<\/p>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 470.25pt\">\n<caption>Figure 15.21: Table showing taphonomic processes that affect the preservation of bones. A. Rodent gnawing. B. Carnivore damage. C. Burned bone. D. Root etching. E. Weathering. F. Cut marks. Credit: A. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Rodent gnawing (Figure 15.26)<\/a>, B. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Carnivore damage (Figure 15.27)<\/a>, C. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Burned bone (Figure 15.28)<\/a>, D. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Root etching (Figure 15.29)<\/a>, E. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Weathering (Figure 15.30)<\/a>, and F. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Cut marks (Figure 15.30)<\/a>, all original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone are under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-NC 4.0 License<\/a>.<\/caption>\n<thead>\n<tr style=\"height: 52.5pt\">\n<td class=\"Table1-C\" style=\"padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Taphonomic Process<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 1pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Definition<\/p>\n<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table1-R\" style=\"height: 190.5pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Rodent Gnawing<\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: center\"><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image19-2.png\" alt=\"Parallel tooth marks etched by a rodent\u2019s front teeth visible on the end of an animal bone.\" width=\"564\" height=\"422\" \/><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">When rodents, such as rats and mice, chew on bone, they leave sets of parallel grooves. The shallow grooves are etched by the rodent\u2019s incisors.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 166.75pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Carnivore Damage<\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: center\"><strong><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image23-4.png\" alt=\"Pit marks from the canines of a carnivore visible on the surface of an animal bone.\" width=\"410\" height=\"272\" \/><\/strong><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Carnivores may leave destructive dental marks on bone. The tooth marks may be visible as pit marks or punctures from the canines, as well as extensive gnawing or chewing of the ends of the bones to retrieve marrow.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 177pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Burned Bone<\/strong><\/p>\n<p class=\"import-Normal\"><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image20-5.png\" alt=\"Burned animal bone fragments pictured at different stages of thermal damage.\" width=\"512\" height=\"342\" \/><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Fire causes observable damage to bone. Temperature and the amount of time bone is heated affect the appearance of the bone. Very high temperatures can crack bone and result in white coloration. Color gradients are visible in between high and lower temperatures, with lower temperatures resulting in black coloration from charring. Cracking can also reveal information about the directionality of the burn.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 169.75pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Root Etching<\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: center\"><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image15-4.png\" alt=\"Animal bone with prominent, discolored grooves where roots leached nutrients from bone\u2019s surface.\" width=\"512\" height=\"342\" \/><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Plant roots can etch the outer surface of bone, leaving grooves where the roots attached as they leached nutrients. During this process, the plant\u2019s roots secrete acid that breaks down the surface of the bone.<\/p>\n<p class=\"import-Normal\">\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 170.5pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Weathering<\/strong><\/p>\n<p class=\"import-Normal\"><strong><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image9.png\" alt=\"Cracking and exfoliation of the surface of an animal bone. \" width=\"512\" height=\"342\" \/><\/strong><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Many different environmental conditions affect bone. River transport can smooth the surface of the bone due to water abrasion. Sunlight can bleach the exposed surface of bone. Dry and wet environments or the mixture of both types of environments can cause cracking and exfoliation of the surface. Burial in different types of soil can cause discoloration, and exposure can cause degreasing.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 169.75pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Cut Marks<\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: left\"><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image8-2.png\" alt=\"Thin vertical lines and cuts are visible along the bone.\" width=\"512\" height=\"342\" \/><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Humans may alter bone by cutting, scraping, or sawing it directly or in the process of removing tissue. The groove pattern\u2014that is, the depth and width of the cuts\u2014can help identify the tool used in the cutting process.<\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<h2 class=\"import-Normal\">Ethics and Human Rights<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Working with human remains requires a great deal of consideration and respect for the dead. Forensic anthropologists have to think about the ethics of our use of human remains for scientific purposes. How do we conduct casework in the most respectable manner possible? While there are a wide range of ethical considerations to consider when contemplating a career in forensic anthropology, this chapter will focus on two major categories: working with human remains and acting as an expert within the medicolegal system.<\/p>\n<h3 class=\"import-Normal\"><strong>Working with Human Remains<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropologists work with human remains in a number of contexts, including casework, excavation, research, and teaching. When working with human remains, it is always important to use proper handling techniques. To prevent damage to skeletal remains, bones should be handled over padded surfaces. Skulls should never be picked up by placing fingers in the eye orbits, foramen magnum (hole at the base of the skull for entry of the spinal cord), or through the zygomatic arches (cheekbones). Human remains, whether related to casework, fieldwork, donated skeletal collections, or research, were once living human beings. It is important to always bear in mind that work with remains should be ingrained with respect for the individual and their relatives. In addition to fieldwork, casework, and teaching, anthropologists are often invited to work with remains that come from a bioarchaeological context or from a human rights violation. While this discussion of ethics is not comprehensive, two case examples will be provided below in which an anthropologist must consider the ethical standards outlined above.<\/p>\n<h3 class=\"import-Normal\"><strong>Modern Human Rights Violations<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropologists may also be called to participate in criminal investigations involving human rights violations. Anthropological investigations may include assistance with identifications, determination of the number of victims, and trauma analyses. In this role, forensic anthropologists play an integral part in promoting human rights, preventing future human rights violations, and providing the evidence necessary to prosecute those responsible for past events. A few ethical considerations for the forensic anthropologist involved in human rights violations include the use of appropriate standards of identification, presenting reliable and unbiased testimony, and maintaining preservation of evidence. For a more comprehensive history of forensic anthropological contributions to human rights violations investigations, see Ubelaker 2018.<\/p>\n<h3 class=\"import-Normal\"><strong>Acting as an Expert in the Medicolegal System<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In addition to the ethical considerations involved in working with human skeletal remains, forensic anthropologists must abide by ethical standards when they act as experts within the medicolegal system. The role of the forensic anthropologist within the medicolegal system is primarily to provide information to the medical examiner or coroner that will aid in the identification process or determination of cause and manner of death. Forensic anthropologists also may be called to testify in a court of law. In this capacity, forensic anthropologists should always abide by a series of ethical guidelines that pertain to their interpretation, presentation, and preservation of evidence used in criminal investigations. First and foremost, practitioners should never misrepresent their training or education. When appropriate, outside opinions and assistance in casework should be requested (e.g., consulting a radiologist for radiological examinations or odontologist for dental exams). The best interest of the decedent should always take precedence. All casework should be conducted in an unbiased way, and financial compensation should never be accepted as it can act as an incentive to take a biased stance regarding casework. All anthropological findings should be kept confidential, and release of information is best done by the medical examiner or coroner. Finally, while upholding personal ethical standards, forensic anthropologists are also expected to report any perceived ethical violations committed by their peers.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ethical standards for the field of forensic anthropology are outlined by the Organization of Scientific Area Committees (OSAC) for Forensic Science, administered by the National Institute of Standards and Technology (NIST). OSAC and NIST recently began an initiative to develop standards that would strengthen the practice of forensic science both in the United States and internationally. OSAC\u2019s main objective is to \u201cstrengthen the nation\u2019s use of forensic science by facilitating the development of technically sound forensic science standards and by promoting the adoption of those standards by the forensic science community\u201d (NIST n.d.). Additionally, OSAC promotes the establishment of best practices and other guidelines to ensure that forensic science findings and their presentation are reliable and reproducible (NIST 2023).<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Native American Graves Protection and Repatriation Act (NAGPRA)<\/h2>\n<p class=\"import-Normal\">There is a long history in the <span style=\"background-color: #00ffff\">United States<\/span> of systematic disenfranchisement of Native American people, including lack of respect for tribal sovereignty. This includes the egregious treatment of Native American human remains. Over several centuries, thousands of Native American remains were removed from tribal lands and held at institutions in the United States, such as museums and universities.<\/p>\n<p class=\"import-Normal\">In 1990, a landmark human rights federal law, the Native American Graves Protection and Repatriation Act (NAGPRA), spurred change in the professional standards and practice of biological anthropology and archaeology. NAGPRA established a legal avenue to provide protection for and repatriation of Native American remains, cultural items, and sacred objects removed from Federal or tribal lands to Native American lineal descendants, Indian tribes, and Native Hawaiian organizations. Human remains and associated artifacts, curated in museum collections and federally funded institutions, are subject to three primary provisions outlined by the NAGPRA statute: (1) protection for Native graves on federal and private land; (2) recognition of tribal authority on such lands; and (3) the requirement that all Native skeletal remains and associated artifacts be inventoried and culturally affiliated groups be consulted concerning decisions related to ownership and final disposition (Rose, Green, and Green 1996). NAGPRA legislation was enacted to ensure ethical consideration and treatment of Native remains and to improve dialogue between scientists and Native groups.<\/p>\n<ul>\n<li>For more information about NAGPRA, visit the <a href=\"https:\/\/www.usbr.gov\/nagpra\/\" target=\"_blank\" rel=\"noopener\">Bureau of Reclamation NAGPRA website<\/a><\/li>\n<li>To read the text of the law, visit the <a href=\"https:\/\/www.congress.gov\/bill\/101st-congress\/house-bill\/5237\">US Congress NAGPRA law website<\/a>.<\/li>\n<li>For further discussion of NAGPRA history, please see <a href=\"https:\/\/textbooks.whatcom.edu\/tracesarchaeology\/\" target=\"_blank\" rel=\"noopener\"><em>TRACES: <\/em><em>An Open Invitation to <\/em><em>Archaeology <\/em>open textbook website<\/a><em><br \/>\n<\/em><\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Becoming a Forensic Anthropologist<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"background-color: #ff99cc\">What does it take to be a forensic anthropologist? Forensic anthropologists are first and foremost anthropologists. While many forensic anthropologists have an undergraduate degree in anthropology, they may also major in biology, criminal justice, pre-law, pre-med, and many other related fields. Practicing forensic anthropologists typically have an advanced degree, either a Master\u2019s or Doctoral degree in Anthropology. Additional training and experience in archaeology, the medico-legal system, rules of evidence, and expert witness testimony are also common. Practicing forensic anthropologists are also encouraged to be board-certified through the American Board of Forensic Anthropology (ABFA). Learn more about the field and educational opportunities on the ABFA website: <a class=\"rId111\" style=\"background-color: #ff99cc\" href=\"https:\/\/www.theabfa.org\/coursework\">https:\/\/www.theabfa.org\/coursework<\/a>.<\/span><\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Review Questions<\/h2>\n<ul>\n<li>What is forensic anthropology? What are the seven primary steps involved in a skeletal analysis?<\/li>\n<li>What are the major components of a biological profile? Why are forensic anthropologists often-tasked with creating biological profiles for unknown individuals?<\/li>\n<li>What are the four major types of skeletal trauma?<\/li>\n<li>What is taphonomy, and why is an understanding of taphonomy often critical in forensic anthropology analyses?<\/li>\n<li>What are some of the ethical considerations faced by forensic anthropologists?<\/li>\n<\/ul>\n<\/div>\n<h2>About the Authors<\/h2>\n<p><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image24-4.jpg\" alt=\"A woman with straight blonde hair smiles at the camera. \" width=\"191\" height=\"254\" \/><\/p>\n<h3 class=\"import-Normal\"><strong>Ashley Kendell, Ph.D.<\/strong><\/h3>\n<p class=\"import-Normal\">California State University, Chico, <a class=\"rId113\" href=\"mailto:akendell@csuchico.edu\">akendell@csuchico.edu<\/a><\/p>\n<p class=\"import-Normal\">Dr. Ashley Kendell is currently an associate professor and forensic anthropologist at Chico State. Prior to beginning her position at Chico State, she was a visiting professor at the University of Montana and the forensic anthropologist for the state of Montana. Dr. Kendell obtained her doctorate from Michigan State University, and her research interests include skeletal trauma analysis and digitization and curation methods for digital osteological data. She is also a Registry Diplomate of the American Board of Medicolegal Death Investigators. Throughout her doctoral program, she worked as a medicolegal death investigator for the greater Lansing, Michigan, area and was involved in the investigation of over 200 forensic cases.<\/p>\n<p class=\"import-Normal\"><strong><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image4.jpg\" alt=\"A woman with straight brown hair pulled back smiles at the camera. \" width=\"194\" height=\"258\" \/><\/strong><\/p>\n<h3 class=\"import-Normal\"><strong>Alex Perrone, M.A., M.S.N, R.N., P.H.N.<\/strong><\/h3>\n<p class=\"import-Normal\">Butte Community College, <a class=\"rId115\" href=\"mailto:perroneal@butte.edu\">perroneal@butte.edu<\/a><\/p>\n<p class=\"import-Normal\">Alex Perrone is a lecturer in anthropology at Butte Community College. She is also a Registered Nurse and a certified Public Health Nurse. She is a former Supervisor of the Human Identification Laboratory in the Department of Anthropology at California State University, Chico. Her research interests include bioarchaeology, paleopathology, forensic anthropology, skeletal biology, California prehistory, and public health. She has worked on bioarchaeological and archaeological projects in Antigua, California, Hawaii, Greece, and the UK, and was an archaeological technician for the USDA Forest Service. She assisted with training courses for local and federal law enforcement agencies and assisted law enforcement agencies with the recovery and analysis of human remains.<\/p>\n<p class=\"import-Normal\" data-wp-editing=\"1\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image1-1.jpg\" alt=\"A woman with curly brown, shoulder-length hair smiles at the camera.\" width=\"190\" height=\"253\" \/><\/p>\n<h3 class=\"import-Normal\"><strong>Colleen Milligan, Ph.D.<\/strong><\/h3>\n<p class=\"import-Normal\">California State University, Chico, <a class=\"rId117\" href=\"mailto:cfmilligan@csuchico.edu\">cfmilligan@csuchico.edu<\/a><\/p>\n<p class=\"import-Normal\">Dr. Colleen Milligan is a biological and forensic anthropologist with research interests in bioarchaeology, skeletal biology, and forensic anthropology. She has been a Fellow with the Department of Homeland Security and has assisted in forensic anthropology casework and recoveries in the State of Michigan and California. She has also assisted in community outreach programs in forensic anthropology and forensic science, as well as recovery training courses for local, state, and federal law enforcement officers. She is a certified instructor through Peace Officers Standards and Training (POST). Dr. Milligan serves as the current co-director of the Chico State Human Identification Laboratory.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration<\/h2>\n<p><a href=\"https:\/\/www.theabfa.org\/coursework\" target=\"_blank\" rel=\"noopener\">The American Board of Forensic Anthropology (ABFA)<\/a><\/p>\n<p><a href=\"https:\/\/www.aafs.org\/\" target=\"_blank\" rel=\"noopener\">The American Academy of Forensic Sciences (AAFS)<\/a><\/p>\n<p><a href=\"https:\/\/www.nist.gov\/organization-scientific-area-committees-forensic-science\" target=\"_blank\" rel=\"noopener\">The Organization of Scientific Area Committees for Forensic Science (OSAC)<\/a><\/p>\n<p><a href=\"https:\/\/textbooks.whatcom.edu\/tracesarchaeology\/\" target=\"_blank\" rel=\"noopener\">TRACES Bioarchaeology<\/a><\/p>\n<p><a href=\"https:\/\/transdoetaskforce.org\/\" target=\"_blank\" rel=\"noopener\">Trans Doe Task Force<\/a><\/p>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Adams, Bradley J., and Lyle W. Konigsberg, eds. 2008. <em>Recovery, Analysis, and Identification of Commingled Remains<\/em>. Totowa, NJ: Humana Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Beatrice, Jared S., and Angela Soler. 2016. \u201cSkeletal Indicators of Stress: A Component of the Biocultural Profile of Undocumented Migrants in Southern Arizona.\u201d <em>Journal of Forensic Sciences <\/em>61 (5): 1164\u20131172.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Berg, Gregory E. 2017. \u201cSex Estimation of Unknown Human Skeletal Remains.\u201d In <em>Forensic Anthropology: A Comprehensive Introduction, Second Edition<\/em>, edited by Natalie R. Langley and MariaTeresa A. Tersigni-Tarrant, 143\u2013159. Boca Raton, FL: CRC Press.<\/p>\n<p class=\"import-Normal\">Bethard, Jonathan D., and Elizabeth A. DiGangi. 2020. \u201cLetter to the Editor\u2014Moving Beyond a Lost Cause: Forensic Anthropology and Ancestry Estimates in the United States.\u201d <em>Journal of Forensic Sciences<\/em> 65 (5): 1791\u20131792.<\/p>\n<p class=\"import-Normal\">Birkby, Walter H., Todd W. Fenton, and Bruce E. Anderson. 2008. \u201cIdentifying Southwest Hispanics Using Nonmetric Traits and the Cultural Profile.\u201d <em>Journal of Forensic Sciences <\/em>53 (1): 29\u201333.<\/p>\n<p class=\"import-Normal\">Blatt, Samantha, Amy Michael, and Lisa Bright. Forthcoming. \u201cBioarchaeology: Interpreting Human Behavior from Skeletal Remains.\u201d In <em>TRACES: <\/em><em>An Open Invitation to <\/em><em>Archaeology<\/em>. https:\/\/textbooks.whatcom.edu\/tracesarchaeology\/.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Brooks, S., and J. M. Suchey. 1990. \u201cSkeletal Age Determination Based on the Os Pubis: A Comparison of the Acs\u00e1di-Nemesk\u00e9ri and Suchey-Brooks Methods.\u201d <em>Human Evolution <\/em>5 (3): 227\u2013238.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Buchanan, Shelby. 2014. \u201cBone Modification in Male to Female Transgender Surgeries: Considerations for the Forensic Anthropologist.\u201d MA thesis, Department of Geography and Anthropology, Louisiana State University, Baton Rouge.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Cunningham, Craig, Louise Scheuer, and Sue Black. 2016. <em>Developmental Juvenile Osteology, Second Edition<\/em>. London: Elsevier Academic Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Galloway, Alison, ed. 1999. <em>Broken Bones: Anthropological Analysis of Blunt Force Trauma<\/em>. Springfield, IL: Charles C. Thomas Publisher, LTD.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Hefner, Joseph T., and Kandus C. Linde. 2018. <em>Atlas of Human Cranial <\/em><em>Macromorphoscopic<\/em><em> Traits<\/em>. San Diego: Academic Press.<\/p>\n<p class=\"import-Normal\">\u0130\u015fcan, M. Y., S. R. Loth, and R. K. Wright. 1984. \u201cAge Estimation from the Rib by Phase Analysis: White Males.\u201d <em>Journal of Forensic Sciences <\/em>29 (4): 1094\u20131104.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">\u0130\u015fcan, M. Y., S. R. Loth, and R. K. Wright. 1985. \u201cAge Estimation from the Rib by Phase Analysis: White Females.\u201d <em>Journal of Forensic Sciences <\/em>30 (3): 853\u2013863.Katz, Darryl, and Judy Myers Suchey. 1986. \u201cAge Determination of the Male Os Pubis.\u201d <em>American Journal of Physical Anthropology <\/em>69 (4): 427\u2013435.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Komar, Debra A., and Jane E. Buikstra. 2008. <em>Forensic Anthropology: Contemporary Theory and Practice<\/em>. New York: Oxford University Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Langley, Natalie R., Alice F. Gooding, and MariaTeresa Tersigni-Tarrant. 2017. \u201cAge Estimation Methods.\u201d In <em>Forensic Anthropology: A Comprehensive Introduction, Second Edition<\/em>, edited by Natalie R. Langley and MariaTeresa A. Tersigni-Tarrant, 175\u2013191. Boca Raton, FL: CRC Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Lovell, Nancy C. 1997. \u201cTrauma Analysis in Paleopathology.\u201d <em>Yearbook of Physical Anthropology<\/em> 104 (S25): 139\u2013170.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Native American Graves Protection and Repatriation Act (NAGPRA) 1990 (25 U.S. Code 3001 et seq.)<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">NIST (National Institute of Standards and Technology). N.d. \u201cThe Organization of Scientific Area Committees for Forensic Science.\u201d Accessed April 18, 2023. <a class=\"rId120\" href=\"https:\/\/www.nist.gov\/topics\/organization-scientific-area-committees-forensic-science\">https:\/\/www.nist.gov\/topics\/organization-scientific-area-committees-forensic-science<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ousley, Stephen. 1995. \u201cShould We Estimate Biological or Forensic Stature?\u201d <em>Journal of Forensic Sciences<\/em> 40(5): 768\u2013773.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Phenice, T. W. 1969. \u201cA Newly Developed Visual Method of Sexing the Os Pubis.\u201d <em>American Journal of Physical Anthropology<\/em> 30 (2): 297\u2013302.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Rose, Jerome C., Thomas J. Green, and Victoria D. Green. 1996. \u201cNAGPRA Is Forever: Osteology and the Repatriation of Skeletons.\u201d <em>Annual Review of Anthropology <\/em>25: 81\u2013103.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Schaefer, Maureen, Sue Black, and Louise Scheuer. <em>Juvenile Osteology: A Laboratory and Field Manua<\/em>l. 2009. San Diego: Elsevier Academic Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Schall, Jenna L., Tracy L. Rogers, and Jordan D. Deschamps-Braly. 2020. \u201cBreaking the Binary: The Identification of Trans-women in Forensic Anthropology.\u201d <em>Forensic Science International<\/em> 309: 110220. https:\/\/doi.org\/10.1016\/j.forsciint.2020.110220.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2010a. \u201cPersonal Identification.\u201d Last modified June 3, 2010. <a class=\"rId121\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_personal_identification.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_personal_identification.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2010b. \u201cSex Assessment.\u201d Last modified June 3, 2010. <a class=\"rId122\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_sex_assessment.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_sex_assessment.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2011. \u201cTrauma Analysis.\u201d Last modified May 27, 2011. <a class=\"rId123\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_trauma.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_trauma.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2012. \u201cStature Estimation.\u201d Last modified August 2, 2012. <a class=\"rId124\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_stature_estimation.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_stature_estimation.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2013. \u201cAge Estimation.\u201d Last modified January 22, 2013. <a class=\"rId125\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_age_estimation.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_age_estimation.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Soler, Angela, and Jared S. Beatrice. 2018. \u201cExpanding the Role of Forensic Anthropology in Humanitarian Crisis: An Example from the USA-Mexico Border. In <em>Sociopolitics of Migrant Death and Repatriation: Perspectives from Forensic Science<\/em>, edited by Krista E. Latham and Alyson J. O\u2019Daniel, 115\u2013128. New York: Springer.<\/p>\n<p class=\"import-Normal\">Soler, Angela, Robin Reineke, Jared Beatrice, and Bruce E. Anderson. 2019. \u201cEtched in Bone: Embodied Suffering in the Remains of Undocumented Migrants.\u201d <em>In<\/em> <em>The Border and Its Bodies: The Embodiment of Risk along the U.S.-M\u00e9xico Line<\/em>, edited by Thomas E. Sheridan and Randall H. McGuire, 173\u2013207. Tucson: University of Arizona Press.<\/p>\n<p class=\"import-Normal\">Stull, Kyra E., Eric J. Bartelink, Alexandra R. Klales, Gregory E. Berg, Michael W. Kenyhercz, Erica N. L\u2019Abb\u00e9, Matthew C. Go, et al.. 2021. \u201cCommentary on: Bethard JD, DiGangi EA. Letter to the Editor\u2014Moving Beyond a Lost Cause: Forensic Anthropology and Ancestry Estimates in the United States. J Forensic Sci. 2020;65(5):1791\u20132. doi: 10.1111\/1556-4029.14513.\u201d <em>Journal of Forensic Sciences <\/em>66 (1): 417\u2013420.<\/p>\n<p class=\"import-Normal\">Tallman, Sean D., Caroline D. Kincer, and Eric D. Plemons. 2022. \u201cCentering Transgender Individuals in Forensic Anthropology and Expanding Binary Sex Estimation in Casework and Research.\u201d Special issue, \u201cDiversity and Inclusion,\u201d <em>Forensic Anthropology<\/em> 5 (2): 161\u2013180.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Tersigni-Tarrant, MariaTeresa A., and Natalie R. Langley. 2017. \u201cHuman Osteology.\u201d In <em>Forensic Anthropology: A Comprehensive Introduction, Second Edition<\/em>, edited by Natalie R. Langley and MariaTeresa A. Tersigni-Tarrant, 81\u2013109. Boca Raton, FL: CRC Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ubelaker, Douglas H. 2018. \u201cA History of Forensic Anthropology.\u201d Special issue, \u201cCentennial Anniversary Issue of AJPA,\u201d <em>American Journal of Physical Anthropology<\/em> 165 (4): 915\u2013923.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">White, Tim D., and Pieter A. Folkens. 2005. <em>The Human Bone Manual<\/em>. Burlington, MA: Elsevier Academic Press.<\/p>\n<p class=\"import-Normal\">Winburn, Allysha P., and Bridget Algee-Hewitt. 2021. \u201cEvaluating Population Affinity Estimates in Forensic Anthropology: Insights from the Forensic Anthropology Database for Assessing Methods Accuracy (FADAMA).\u201d <em>Journal of Forensic Sciences<\/em> 66 (4): 1210\u20131219.<\/p>\n<p class=\"import-Normal\">Winburn, Allysha Powanda, Sarah Kiley Schoff, and Michael W. Warren. 2016. \u201cAssemblages of the Dead: Interpreting the Biocultural and Taphonomic Signature of Afro- Cuban Palo Practice in Florida.\u201d <em>Journal of African Diaspora Archaeology and Heritage <\/em>5 (1): 1\u201337.<\/p>\n<\/div>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1779\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1779\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\">Ashley Kendell, Ph.D., California State University, Chico<\/p>\n<p class=\"import-Normal\">Alex Perrone, M.A., M.S.N, R.N., P.H.N., Butte Community College<\/p>\n<p class=\"import-Normal\">Colleen Milligan, Ph.D., California State University, Chico<\/p>\n<p class=\"import-Normal\"><em>This chapter is a revision from \"<\/em><a class=\"rId7\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\"><em>Chapter 15: Bioarchaeology and Forensic Anthropology<\/em><\/a><em>\u201d by Ashley Kendell, Alex Peronne, and Colleen Milligan. In <\/em><a class=\"rId8\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, first edition<\/em><\/a><em>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under <\/em><a class=\"rId9\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\"><em>CC BY-NC 4.0<\/em><\/a><em>. <\/em><\/p>\n<p class=\"import-Normal\"><strong>Content Warning and Disclaimer:<\/strong> This chapter includes images of human remains as well as discussions centered on human skeletal analyses. All images are derived from casts, sketches, nonhuman skeletal material, as well as non-Indigenous skeletal materials curated within the CSU, Chico Human Identification Lab, and the Hartnett-Fulginiti donated skeletal collection.<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\"><span style=\"color: #ffffff\">Learning Objectives<\/span><\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Define forensic anthropology as a subfield of biological anthropology.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Describe the seven steps carried out during skeletal analysis.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Outline the four major components of the biological profile.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Contrast the four categories of trauma.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Explain how to identify the different taphonomic agents that alter bone.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Discuss ethical considerations for forensic anthropology.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1214\">Forensic anthropology<\/a><\/strong> is a subfield of biological anthropology and an applied area of anthropology. Forensic anthropologists use skeletal analysis to gain information about humans in the present or recent past, then they apply this information within a medicolegal context. This means that forensic anthropologists specifically conduct their analysis on recently deceased individuals (typically within the last 50 years) as part of investigations by law enforcement. Forensic anthropologists can assist law enforcement agencies in several different ways, including aiding in the identification of human remains whether they are complete, fragmentary, burned, scattered, or decomposed. Additionally, forensic anthropologists can help determine what happened to the deceased at or around the time of death as well as what processes acted on the body after death (e.g., whether the remains were scattered by animals, whether they were buried in the ground, or whether they remained on the surface as the soft tissue decomposed).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Many times, because of their expertise in identifying human skeletal remains, forensic anthropologists are called to help with outdoor search-and-recovery efforts, such as locating remains scattered across the surface or carefully excavating and documenting buried remains. In other cases, forensic anthropologists recover remains after natural disasters or accidents, such as fire scenes, and can help identify whether each bone belongs to a human or an animal. Forensic anthropology spans a wide scope of contexts involving the law, including incidences of mass disasters, genocide, and war crimes.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">A point that can be somewhat confusing for students is that although the term <em>forensic<\/em> is included in this subfield of biological anthropology, there are many forensic techniques that are not included in the subfield. Almost exclusively, forensic anthropology deals with skeletal analysis. While this can include the comparison of antemortem (before death) and postmortem (after death) radiographs to identify whether remains belong to a specific person, or using photographic superimposition of the cranium, it does not include analyses beyond the skeleton. For example, blood-spatter analysis, DNA analysis, fingerprints, and material evidence collection do not fall under the scope of forensic anthropology.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">So, what can forensic anthropologists glean from bones alone? Forensic anthropologists can address a number of questions about a human individual based on their skeletal remains. Some of those questions are as follows: How old was the person? Was the person biologically male or female? How tall was the person? What happened to the person at or around their time of death? Were they sick? The information from the skeletal analysis can then be matched with missing persons records, medical records, or dental records, aiding law enforcement agencies with identifications and investigations.<\/p>\n<h2 class=\"import-Normal\">Skeletal Analysis<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropology relies on skeletal analysis to reveal information about the deceased. <span style=\"background-color: #00ffff\">The methodology and approaches outlined below are specific to the United States.<\/span> Forensic anthropological methods differ depending on the country conducting an investigation. In the United States, there are typically seven steps or questions to the process:<\/p>\n<ul>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Is it bone?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Is it human?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Is it modern or archeological?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">How many individuals are present or what is the minimum number of individuals (MNI)?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Who is it?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Is there evidence of trauma before or around the time of death?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">What happened to the remains after death?<\/li>\n<\/ul>\n<h3 class=\"import-Normal\"><strong>Is It Bone?<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">One of the most important steps in any skeletal analysis starts with determining whether or not material suspected to be bone is in fact bone. Though it goes without saying that a forensic anthropologist would only carry out analysis on bone, this step is not always straightforward. Whole bones are relatively easy to identify, but determining whether or not something is bone becomes more challenging once it becomes fragmentary. As an example, in high heat such as that seen on fire scenes, bone can break into pieces. During a house fire with fatalities, firefighters watered down the burning home. After the fire was extinguished, the sheetrock (used to construct the walls of the home) was drenched and crumbled. The crumbled sheetrock was similar in color and form to burned, fragmented bone, therefore mistakable for human remains (Figure 15.1). Forensic anthropologists on scene were able to separate the bones from the construction material, helping to confirm the presence of bone and hence the presence of individual victims of the fire. In this case, forensic anthropologists were able to recognize the anatomical and layered structure of bone and were able to distinguish it from the uniform and unlayered structure of sheetrock.<\/p>\n<p class=\"import-Normal\"><strong><img class=\"aligncenter\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/06\/image3.png\" alt=\"Long rectangular sheetrock with exposed porous surface.\" width=\"182\" height=\"208\" \/><\/strong><\/p>\n<figure style=\"width: 372px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image6-1.png\" alt=\"Two examples of sheetrock with dried or burnt surfaces.\" width=\"372\" height=\"210\" \/><figcaption class=\"wp-caption-text\">Figure 15.1: Burned sheetrock used as building material appears similar to human bone but can be differentiated by the fact that it is the same density throughout. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Example of burned sheetrock (Figure 15.1)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">As demonstrated by the example above, both the macrostructure (visible with the naked eye) and microstructure (visible with a microscope) of bone are helpful in bone identification. Bones are organs in the body made up of connective tissue. The connective tissue is hardened by a mineral deposition, which is why bone is rigid in comparison to other connective tissues such as cartilage (Tersigni-Tarrant and Langley 2017, 82\u201383; White and Folkens 2005, 31). In a living body, the mineralized tissue does not make up the only component of bone\u2014there are also blood, bone marrow, cartilage, and other types of tissues. However, in dry bone, two distinct layers of the bone are the most helpful for identification. The outer layer is made up of densely arranged osseous (bone) tissue called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1216\">compact (cortical) bone<\/a><\/strong>. The inner layer is composed of much more loosely organized, porous bone tissue whose appearance resembles that of a sponge, hence the name <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1218\"><strong>spongy (trabecular) bone<\/strong><\/a>. Knowing that most bone contains both layers helps with the macroscopic identification of bone (Figures 15.2, 15.3). For example, a piece of coconut shell might look a lot like a fragment of a human skull bone. However, closer inspection will demonstrate that coconut shell only has one very dense layer, while bone has both the compact and spongy layers.<\/p>\n<figure style=\"width: 380px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image27-1.png\" alt=\"Drawing showing thick exterior compact bone and porous internal cortical bone.\" width=\"380\" height=\"371\" \/><figcaption class=\"wp-caption-text\">Figure 15.2: Cross section of human long bone with compact and cortical bone layers visible. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Cross section of human long bone (Figure 15.2)<\/a> original to<a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"> Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p>&nbsp;<\/p>\n<figure style=\"width: 364px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image25-2.png\" alt=\"Cranial bone cross section called a periosteum with spongy bone (diploe) and compact bone labeled. Compact bone is a thin slice at the top and bottom and is smooth and hard. Spongy bone is in the middle and has irregular holes and indentations throughout. \" width=\"364\" height=\"184\" \/><figcaption class=\"wp-caption-text\">Figure 15.3: Cranial anatomy is slightly different as compared to that of a long bone in cross section. The compact (cortical) bone layers sandwich the spongy (trabecular) bone. One layer of compact bone forms the very outer surface of the skull and the other lines the internal surface of the skull. Credit: <a href=\"https:\/\/cnx.org\/contents\/FPtK1zmh@6.27:kwbeYj9S@3\/Bone-Structure\">Anatomy of a Flat Bone (Anatomy &amp; Physiology, Figure 6.3.3)<\/a> by<a href=\"https:\/\/openstax.org\/\"> OpenStax<\/a> is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/\"> CC BY 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The microscopic identification of bone relies on knowledge of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1220\">osteons<\/a><\/strong>, or bone cells (Figure 15.4). Under magnification, bone cells are visible in the outer, compact layer of bone. The bone cells are arranged in a concentric pattern around blood vessels for blood supply. The specific shape of the cells can help differentiate, for example, a small piece of PVC (white plastic) pipe from a human bone fragment (Figure 15.5).<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 340px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image14-3.png\" alt=\"Microscope image showing clustered osteons. Each has many rings and a dark center.\" width=\"340\" height=\"218\" \/><figcaption class=\"wp-caption-text\">Figure 15.4: Bone microstructure (osteons). Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Bone_(248_12)_Bone_cross_section.jpg\">Bone (248 12) Bone cross section<\/a> by <a href=\"https:\/\/cs.wikipedia.org\/wiki\/Josef_Reischig\">Doc. RNDr. Josef Reischig, CSc.<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p>&nbsp;<\/p>\n<figure style=\"width: 332px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image7-1.png\" alt=\"Flat, white section of PVC. Edges are broken and surface rough.\" width=\"332\" height=\"268\" \/><figcaption class=\"wp-caption-text\">Figure 15.5: Fragments of plastic PVC pipe, such as those seen in this photo, may be mistaken for human bone. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Example of PVC pipe<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Is It Human?<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Once it has been determined that an object is bone, the next logical step is to identify whether the bone belongs to a human or an animal. Forensic anthropologists are faced with this question in everyday practice because human versus nonhuman bone identification is one of the most frequent requests they receive from law enforcement agencies.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">There are many different ways to distinguish human versus nonhuman bone. The morphology (the shape\/form) of human bone is a good place for students to start. Identifying the 206 bones in the adult human skeleton and each bone\u2019s distinguishing features (muscle attachment sites, openings and grooves for nerves and blood vessels, etc.) is fundamental to skeletal analysis.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Nevertheless, there are many animal bones and human bones that look similar. For example, the declawed skeleton of a bear paw looks a lot like a human hand, pig molars appear similar to human molars, and some smaller animal bones might be mistaken for those of an infant. To add to the confusion, fragmentary bone may be even more difficult to identify as human or nonhuman. However, several major differences between human and nonhuman vertebrate bone help distinguish the two.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropologists pay special attention to the density of the outer, compact layer of bone in both the cranium and in the long bones. Human cranial bone has three distinctive layers. The spongy bone is sandwiched between the outer (ectocranial) and inner (endocranial) compact layers. In most other mammals, the distinction between the spongy and compact layers is not always so definite. Secondly, the compact layer in nonhuman mammal long bones can be much thicker than observed in human bone. Due to the increased density of the compact layer, nonhuman bone tends to be heavier than human bone (Figure 15.6).<\/p>\n<figure style=\"width: 399px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image11-2.png\" alt=\"Ring-like cross section of bone.\" width=\"399\" height=\"266\" \/><figcaption class=\"wp-caption-text\">Figure 15.6: The compact layer of this animal bone is very thick, with almost no spongy bone visible. Compare with Figure 15.2 to visualize the difference in structure between human and nonhuman bone. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Animal bone cross section (Figure 15.6)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The size of a bone can also help determine whether it belongs to a human. Adult human bones are larger than subadult or infant bones. However, another major difference between human adult bones and those of a young individual or infant human can be attributed to development and growth of the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1222\">epiphyses<\/a><\/strong> (ends of the bone). The epiphyses of human subadult bones are not fused to the shaft (Figure 15.7). Therefore, if a bone is small and it is suspected to belong to a human subadult or infant, the epiphyses would not be fused. Many small animal bones appear very similar in form compared to adult human bones, but they are much too small to belong to an adult human. Yet they can be eliminated as subadult or infant bones if the epiphyses are fused to the shaft.<\/p>\n<figure style=\"width: 288px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image13-3.png\" alt=\"X-ray image of child\u2019s ankle.\" width=\"288\" height=\"412\" \/><figcaption class=\"wp-caption-text\">Figure 15.7: An x-ray of a subadult\u2019s ankle with the epiphyses of the tibia and fibula visible. The gap between the shaft of the bone and the end of the bone (epiphysis) is the location of the growth plate. Therefore, the growth plate gap is what separates the shafts from the epiphyses in the image. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Tib_fib_growth_plates.jpg\">Tib fib growth plates<\/a> by <a href=\"https:\/\/en.wikipedia.org\/wiki\/User:Gilo1969\">Gilo1969<\/a> at <a href=\"https:\/\/en.wikipedia.org\/wiki\/\">English Wikipedia<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/3.0\/legalcode\">CC BY 3.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Is It Modern or Archaeological? <\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropologists work with modern cases that fall within the scope of law enforcement investigations. Accordingly, it is important to determine whether discovered human remains are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1224\">archaeological<\/a> <\/strong>or forensic in nature. Human remains that are historic are considered archeaological. The scientific study of human remains from archaeological sites is called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1226\">bioarchaeology<\/a><\/strong>.<\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Dig Deeper: Bioarchaeology<\/h2>\n<p class=\"import-Normal\">For readers who are interested in the sister subfield of bioarchaeology, which studies human remains and material culture from the past, please refer to chapter 8 of <em>Bioarchaeology: Interpreting Human Behavior from Skeletal Remains,<\/em> in <em>TRACES: An Open Invitation to Archaeology<\/em> (Blatt, Michael, and Bright forthcoming).<\/p>\n<\/div>\n<p>A forensic anthropologist should begin their analysis by reviewing the context in which the remains were discovered. This will help them understand a great deal about the remains, including determining whether they are archaeological or forensic in nature as well as considering legal and ethical issues associated with the collection, analysis, and storage of human remains (see \u201cEthics and Human Rights\u201d section of this chapter for more information).<\/p>\n<figure style=\"width: 403px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image10-3.png\" alt=\"Four teeth in a person\u2019s mouth. First molar with silver filling.\" width=\"403\" height=\"303\" \/><figcaption class=\"wp-caption-text\">Figure 15.8: A human tooth with a filling. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Filling.jpg#filehistory\">Filling<\/a> by Kauzio has been designated to the <a href=\"https:\/\/creativecommons.org\/share-your-work\/public-domain\/cc0\/\">public domain (CC0)<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The \u201ccontext\u201d refers to the relationship the remains have to the immediate area in which they were found. This includes the specific place where the remains were found, the soil or other organic matter immediately surrounding the remains, and any other objects or artifacts in close proximity to the body. For example, imagine that a set of remains has been located during a house renovation. The remains are discovered below the foundation. Do the remains belong to a murder victim? Or was the house built on top of an ancient burial ground? Observing information from the surroundings can help determine whether the remains are archaeological or modern. How long ago was the foundation of the house erected? Are there artifacts in close proximity to the body, such as clothing or stone tools? These are questions about the surroundings that will help determine the relative age of the remains.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Clues directly from the skeleton may also indicate whether the remains are archaeological or modern. For example, tooth fillings can suggest that the individual was alive recently (Figure 15.8). In fact, filling material has changed over the decades, so the specific type of material used to fix a cavity can be matched with specific time periods. Gold was used in dental work in the past, but more recently composite (a mixture of plastic and fine glass) fillings have become more common.<\/p>\n<h3><strong>How <\/strong><strong>Many Individuals Are Present?<\/strong><\/h3>\n<h4 class=\"import-Normal\"><em>What Is MNI?<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Another assessment that an anthropologist can perform is the calculation of the number of individuals in a mixed burial assemblage. Because not all burials consist of a single individual, it is important to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1268\">burial assemblage<\/a><\/strong> be able to estimate the number of individuals in a forensic context. Quantification of the number of individuals in a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1524\">burial assemblage<\/a><\/strong> can be done through the application of a number of methods, including the following: the Minimum Number of Individuals (MNI), the Most Likely Number of Individuals (MLNI), and the Lincoln Index (LI). The most commonly used method in biological anthropology, and the focus of this section, is determination of the MNI.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The MNI presents \u201cthe minimum estimate for the number of individuals that contributed to the sample\u201d (Adams and Konigsberg 2008, 243). Many methods of calculating MNI were originally developed within the field of zooarchaeology for use on calculating the number of individuals in faunal or animal assemblages (Adams and Konigsberg 2008, 241). What MNI calculations provide is a lowest possible count for the total number of individuals contributing to a skeletal assemblage. Traditional methods of calculating MNI include separating a skeletal assemblage into categories according to the individual bone and the side the bone comes from and then taking the highest count per category and assigning that as the minimum number (Figure 15.9).<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 664px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image28-3.png\" alt=\"Many bone portions laying on individual plastic bags on a table.\" width=\"664\" height=\"441\" \/><figcaption class=\"wp-caption-text\">Figure 15.9: Skeletal elements from a commingled faunal assemblage. Credit: Commingled animal remains from Eden-Farson Pre-Contact site in southwest Wyoming by Matt O\u2019Brien original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Why Calculate MNI?<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In a forensic context, the determination of MNI is most applicable in cases of mass graves, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1232\">commingled burials<\/a><\/strong>, and mass fatality incidents. The term <em>commingled<\/em> is applied to any burial assemblage in which individual skeletons are not separated into separate burials. As an example, the authors of this chapter have observed commingling of remains resulting from mass fatality wildfire events. Commingled remains may also be encountered in events such as a plane or vehicle crash. It is important to remember that in any forensic context, MNI should be referenced and an MNI of one should be substantiated by the fact that there was no repetition of elements associated with the case.<\/p>\n<h3 class=\"import-Normal\"><strong>Constructing the Biological Profile<\/strong><\/h3>\n<h4 class=\"import-Normal\"><em>Who Is It?<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">\u201cWho is it?\u201d is one of the first questions that law enforcement officers ask when they are faced with a set of skeletal remains. To answer this question, forensic anthropologists construct a biological profile (White and Folkens 2005, 405). A <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1228\">biological profile<\/a> <\/strong>is an individual\u2019s identifying characteristics, or biological information, which include the following: biological sex, age at death, stature, population affinity, skeletal variation, and evidence of trauma and pathology.<\/p>\n<h4 class=\"import-Normal\"><em>Assessing Biological Sex <\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Assessment of biological sex is often one of the first things considered when establishing a biological profile because several other parts, such as age and stature estimations, rely on an assessment of biological sex to make the calculations more accurate.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Assessment of biological sex focuses on differences in both morphological (form or structure) and metric (measured) traits in individuals. When assessing morphological traits, the skull and the pelvis are the most commonly referenced areas of the skeleton. These differences are related to sexual dimorphism usually varying in the amount of robusticity seen between males and females. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1230\">Robusticity<\/a> <\/strong>deals with strength and size; it is frequently used as a term to describe a large size or thickness. In general, males will show a greater degree of robusticity than females. For example, the length and width of the mastoid process, a bony projection located behind the opening for the ear, is typically larger in males. The mastoid process is an attachment point for muscles of the neck, and this bony projection tends to be wider and longer in males. In general, cranial features tend to be more robust in males (Figure 15.10).<\/p>\n<figure style=\"width: 601px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image26-3.png\" alt=\"Front and side images of a male (left) and female (right) cranium.\" width=\"601\" height=\"632\" \/><figcaption class=\"wp-caption-text\">Figure 15.10: Anterior and lateral view of a male and female cranium. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Anterior and lateral view of a male and female cranium (Figure 15.10)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropo logy<\/a> by Ashley Kendell is a collective work under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Includes <a href=\"https:\/\/boneclones.com\/product\/modern-human-asian-female-skull-BC-149\/category\/all-human-skulls\/human-anatomy\">Human Female Asian Skull<\/a> and <a href=\"https:\/\/boneclones.com\/product\/human-asian-male-skull-BC-016\/category\/all-human-skulls\/human-anatomy\">Human Male Asian Skull<\/a> by <a href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a>, used by permission.]<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">When considering the pelvis, the features associated with the ability to give birth help distinguish females from males. During puberty, estrogen causes a widening of the female pelvis to allow for the passage of a baby. Several studies have identified specific features or bony landmarks associated with the widening of the hips, and this section will discuss one such method. The Phenice Method (Phenice 1969) is traditionally the most common reference used to assess morphological characteristics associated with sex. The Phenice Method specifically looks at the presence or absence of (1) a ventral arc, (2) the presence or absence of a subpubic concavity, and (3) the width of the medial aspect of the ischiopubic ramus (Figure 15.11). When present, the ventral arc, a ridge of bone located on the ventral surface of the pubic bone, is indicative of female remains. Likewise the presence of a subpubic concavity and a narrow medial aspect of the ischiopubic ramus is associated with a female sex estimation. Assessments of these features, as well as those of the skull (when both the pelvis and skull are present), are combined for an overall estimation of sex.<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 1603px\" class=\"wp-caption alignnone\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image29-3.png\" alt=\"Male and female os coxae (anterior portions).\" width=\"1603\" height=\"582\" \/><figcaption class=\"wp-caption-text\">Figure 15.11: Features associated with the Phenice Method. Images derived from CSU-HIL donated skeletal collection. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Features associated with the Phenice Method (Figure 15.11)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Colleen Milligan is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Metric analyses are also used in the estimation of sex. Measurements taken from every region of the body can contribute to estimating sex through statistical approaches that assign a predictive value of sex. These approaches can include multiple measurements from several skeletal elements in what is called multivariate (multiple variables) statistics. Other approaches consider a single measurement, such as the diameter of the head of the femur, of a specific element in a univariate (single variable) analysis (Berg 2017, 152\u2013156).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">It is important to note that, although forensic anthropologists usually begin assessment of biological profile with biological sex, there is one major instance in which this is not appropriate. The case of two individuals found in California, on July 8, 1979, is one example that demonstrates the effect age has on the estimation of sex. The identities of the two individuals were unknown; therefore, law enforcement sent them to a lab for identification. A skeletal analysis determined that the remains represented one adolescent male and one adolescent female, both younger than 18 years of age. This information did not match with any known missing children at the time.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In 2015, the cold case was reanalyzed, and DNA samples were extracted. The results indicated that the remains were actually those of two girls who went missing in 1978. The girls were 15 years old and 14 years old at the time of death. It is clear that the 1979 results were incorrect, but this mistake also provides the opportunity to discuss the limitations of assessing sex from a subadult skeleton.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Assessing sex from the human skeleton is based on biological and genetic traits associated with females and males. These traits are linked to differences in sexual dimorphism and reproductive characteristics between females and males. The link to reproductive characteristics means that most indicators of biological sex do not fully manifest in prepubescent individuals, making estimations of sex unreliable in younger individuals (SWGANTH 2010b). This was the case in the example of the 14-year-old girl. When examined in 1979, her remains were misidentified as male because she had not yet fully developed female pelvic traits.<\/p>\n<h4 class=\"import-Normal\"><em>Sex vs. Gender<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Biological sex is a different concept than <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1234\">gender<\/a><\/strong>. While biological anthropologists can estimate sex from the skeleton, estimating an individual\u2019s gender would require a greater context because gender is defined culturally rather than biologically. Take, for example, an individual who identifies as transgender. This individual has a gender identity that is different from their biological sex. The gender identity of any individual depends on factors related to self-identification, situation or context, and cultural factors. <span style=\"background-color: #00ffff\">While in the U.S<\/span>. we have historically thought of sex and gender as binary concepts (male or female), many cultures throughout the world recognize several possible gender identities. In this sense, gender is seen as a continuous or fluid variable rather than a fixed one.<\/p>\n<p class=\"import-Normal\">Historically, forensic anthropologists have used a binary construct to categorize human skeletal remains as either male or female (with the accompanying categories of probable male, probable female, and indeterminate). In the case of transgender and gender nonconforming individuals, the binary approach to sex assessment may delay or hinder identification efforts (Buchanan 2014; Schall, Rogers, and Deschamps-Braly 2020; Tallman, Kincer, and Plemons 2021). As such, many forensic anthropologists have begun to address the inherent problems associated with a binary approach to sex identification and to explore ways of assessing social identity and self-identified gender using skeletal remains and forensic context.<\/p>\n<p class=\"import-Normal\">For the duration of this section, the term <em>transgender<\/em> refers to individuals whose gender identity differs from the sex assigned at birth (Schall, Rogers, and Deschamps-Braly 2020:2). Transgender individuals transition from one gender binary to another, such as male-to-female (MTF) or female-to-male (FTM). While many of the gender-affirming procedures available to trans and gender-nonconforming individuals are focused on soft tissue modifications (e.g., breast augmentation, genital reconstruction, hormone therapies, etc.), there are a number of gender-affirmation surgeries that do leave a permanent record on the skeleton. Generally speaking, FTM transgender people are reported to undergo fewer surgical procedures than do MTF transgender people (Buchanan 2014). The discussion below focuses on Facial Feminization Surgery (FFS), which leaves a permanent record on the human skeleton that may be used to help make an identification.<\/p>\n<p class=\"import-Normal\">FFS refers to a combination of procedures focused on sexually dimorphic features of the face, with the intent of transforming typically male facial features into more feminine forms. Facial Feminization Surgery procedures were developed by Dr. Douglas Ousterhout, a San Francisco based cranio-maxillofacial surgeon, in the mid-1980s (Schall, Rogers, and Deschamps-Braly 2020:2). FFS can include one or a combination of the following: hairline lowering, forehead reduction and contouring, brow lift, reduction rhinoplasty, cheek enhancement, lift lift, lip filling, chin contouring, jaw contouring, and\/or tracheal shave (Buchanan 2014; Schall, Rogers, and Deschamps-Braly 2020:2). Of the procedures outlined previously, four are known to directly affect the facial skeleton: forehead contouring, rhinoplasty, chin contouring, and jaw contouring (Buchanan 2014; Schall, Rogers, and Deschamps-Braly 2020:2).<\/p>\n<p class=\"import-Normal\">Because FFS procedures have been widely documented in the medical (and more recently the forensic anthropological) literature, there are a number of indicators that a forensic anthropologist can use to make more informed evaluations of gender, including evidence of bone remodeling in sexually dimorphic regions of the skull (e.g., forehead, chin, jawline), as well as the presence of plates, pins, or other surgical hardware that may be evidence of FFS (Buchanan 2014; Schall, Rogers, and Deschamps-Braly 2020; Tallman, Kincer, and Plemons 2021). Additionally, some forensic anthropologists suggest cautiously integrating contextual information from the scene, such as personal effects, material evidence, and recovery scene information, into their evaluation of an individual\u2019s social identity (Beatrice and Soler 2016; Birkby, Fenton, and Anderson 2008; Soler and Beatrice 2018; Soler et al. 2019; Tallman, Kincer, and Plemons 2021; Winburn, Schoff, and Warren 2016). The ultimate goal of many skeletal analyses is to make a positive identification on a set of unidentified remains.<\/p>\n<h4 class=\"import-Normal\"><em>Assessment <\/em><em>of Population Affinity<\/em><\/h4>\n<p>In an effort to combat the erroneous assumptions tied to the race concept, forensic anthropologists have attempted to reframe this component of the biological profile. The term <em>race<\/em> is no longer used in casework and teaching. Historically, the word <em>ancestry<\/em> is and was deemed a more appropriate way to describe an individual\u2019s phenotype. However, in more recent years, forensic anthropologists have begun using the term <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1236\">population affinity<\/a><\/strong><em>, <\/em>recognizing that we are basing our analysis on the similarities we see based on the reference samples we have available (Winburn and Algee-Hewitt 2021). An important note here is that it is possible to hinder identifications and harm individuals when tools like estimations of population affinity are misapplied, misinterpreted, or misused. For this reason, the field of forensic anthropology has ongoing conversations about the appropriateness of this analysis in the biological profile (Bethard and DiGangi 2020; Stull et al. 2021).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">We use the term <em>population affinity<\/em> to refer to the variation seen among modern populations\u2014variation that is both genetic and environmentally driven. The word <em>affinity<\/em> refers to similarities or relationships between individuals. As forensic anthropologists, we compare an unknown individual to multiple reference groups and look for the degree of similarity in observable traits with those groups. As noted previously, population affinity can aid law enforcement in their identification of missing persons or unknown skeletal remains.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Within the field of anthropology, the estimation of population affinity has a contentious history, and early attempts at classification were largely based on the erroneous assumption that an individual\u2019s <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1238\">phenotype <\/a><\/strong> (outward appearance) was correlated with their innate intelligence and abilities (see Chapter 13 for a more in-depth discussion of the history of the race concept). The use of the term <em>race<\/em> is deeply embedded in the social context of the United States. In any other organism\/living thing, groups divided according to the biological race concept would be defined as a separate subspecies. The major issue with applying the biological race concept to humans is that there are not enough differences between any two populations to separate on a genetic basis. In other words, <em>biological races do not exist in human populations. <\/em>However, the concept of race has been perpetuated and upheld by sociocultural constructs of race.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The conundrum for forensic anthropologists is the fact that while races do not exist on a biological level, we still socially recognize and categorize individuals based on their phenotype. Clearly, our phenotype is an important factor in not only how we are viewed by others but also how we identify ourselves. It is also a commonly reported variable. Often labeled as \u201crace,\u201d we are asked to report how we self-identify on school applications, government identification, surveys, census reports, and so forth. It follows then that when a person is reported missing, the information commonly collected by law enforcement and sometimes entered into a missing person\u2019s database includes their age, biological sex, stature, and \u201crace.\u201d Therefore, the more information a forensic anthropologist can provide regarding the individual\u2019s physical characteristics, the more he or she can help to narrow the search.<\/p>\n<p class=\"import-Normal\">As an exercise, create a list of all of the women you know who are between the ages of 18 and 24 and approximately 5\u2019 4\u201d to 5\u2019 9\u201d tall. You probably have several dozen people on the list. Now, consider how many females you know who are between the ages of 18 and 24, are approximately 5\u2019 4\u201d to 5\u2019 9\u201d tall, and are Vietnamese. Your list is going to be significantly shorter. That\u2019s how missing persons searches go as well. The more information you can provide regarding a decedent\u2019s phenotype, the fewer possible matches law enforcement are left to investigate. This is why population affinity has historically been included as a part of the biological profile.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Traditionally, population affinity was accomplished through a visual inspection of morphological variants of the skull (morphoscopics). These methods focused on elements of the facial skeleton, including the nose, eyes, and cheek bones. However, in an effort to reduce subjectivity, nonmetric cranial traits are now assessed within a statistical framework to help anthropologists better interpret their distribution among living populations (Hefner and Linde 2018). Based on the observable traits, a macromorphoscopic analysis will allow the practitioner to create a statistical prediction of geographic origin. In essence, forensic anthropologists are using human variation in the estimation of geographic origin, by referencing documented frequencies of nonmetric skeletal indicators or macromorphoscopic traits.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Population affinity is also assessed through metric analyses. The computer program Fordisc is an anthropological tool used to estimate different components of the biological profile, including ancestry, sex, and stature. When using Fordisc, skeletal measurements are input into the computer software, and the program employs multivariate statistical classification methods, including discriminant function analysis, to generate a statistical prediction for the geographic origin of unknown remains based on the comparison of the unknown to the reference samples in the software program. Fordisc also calculates the likelihood of the prediction being correct, as well as how typical the metric data is for the assigned group.<\/p>\n<h4 class=\"import-Normal\"><em>Estimating Age-at-Death<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Estimating age-at-death from the skeleton relies on the measurement of two basic physiological processes: (1) growth and development and (2) degeneration (or aging). From fetal development on, our bones and teeth grow and change at a predictable rate. This provides for relatively accurate age estimates. After our bones and teeth cease to grow and develop, they begin to undergo structural changes, or degeneration, associated with aging. This does not happen at such predictable rates and, therefore, results in less accurate or larger age-range estimations.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">During growth and development stages, two primary methods used for estimations of age of subadults (those under the age of 18) are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1240\">epiphyseal union<\/a><\/strong> and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1244\">dental development.<\/a><\/strong> Epiphyseal union<strong> (<\/strong>or <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1242\">epiphyseal fusion<\/a><\/strong>) refers to the appearance and closure of the epiphyseal plates between the primary centers of growth in a bone and the subsequent centers of growth (see Figure 15.7). Prior to complete union, the cartilaginous area between the primary and secondary centers of growth is also referred to as the growth plates (Schaefer, Black, and Scheuer 2009). Different areas of the skeleton have documented differences in the appearance and closure of epiphyses, making this a reliable method for aging subadult remains (SWGANTH 2013).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">As an example of its utility in the identification process, epiphyseal development was used to identify two subadult victims of a fatal fire in Flint, Michigan, in February 2010. The remains represented two young girls, ages three and four. Due to the intensity of the fire, the subadult victims were differentiated from each other through the appearance of the patella, the kneecap. The patella is a bone that develops within the tendon of the quadriceps muscle at the knee joint. The patella begins to form around three to four years of age (Cunningham, Scheuer, and Black 2016, 407\u2013409). In the example above, radiographs of the knees showed the presence of a patella in the four-year-old girl and the absence of a clearly discernible patella in the three-year-old.<\/p>\n<figure style=\"width: 358px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image18-2.png\" alt=\"Cranial cast of child with exposed maxilla and mandible to see developing dentition.\" width=\"358\" height=\"358\" \/><figcaption class=\"wp-caption-text\">Figure 15.12: Dental development in a subadult. Credit: <a href=\"https:\/\/boneclones.com\/product\/5-year-old-human-child-skull-with-mixed-dentition-exposed-BC-189\">5-year-old Human Child Skull with Mixed Dentition Exposed<\/a> by <a href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a> is used by permission and available here is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Dental development begins during fetal stages of growth and continues until the complete formation and eruption of the adult third molars (if present). The first set of teeth to appear are called deciduous or baby teeth. Individuals develop a total of 20 deciduous teeth, including incisors, canines, and molars. These are generally replaced by adult dentition as an individual grows (Figure 15.12). A total of 32 teeth are represented in the adult dental arcade, including incisors, canines, premolars, and molars. When dental development is used for age estimations, researchers use both tooth-formation patterns and eruption schedules as determining evidence. For example, the crown of the tooth forms first followed by the formation of the tooth root. During development, an individual can exhibit a partially formed crown or a complete crown with a partially formed root. The teeth generally begin the eruption process once the crown of the tooth is complete. The developmental stages of dentition are one of the most reliable and consistent aging methods for subadults (Langley, Gooding, and Tersigni-Tarrant 2017, 176\u2013177).<\/p>\n<figure style=\"width: 403px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image22-3.png\" alt=\"Surfaces of three pubic symphyses: billowy (A) to more flat (B) to rough (C).\" width=\"403\" height=\"224\" \/><figcaption class=\"wp-caption-text\">Figure 15.13: Examples of degenerative changes to the pubic symphysis: (A) young adult; (B) middle adult; (C) old adult. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Example of the progression of degenerative changes to the pubic symphysis (Figure 15.14)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropo logy<\/a> by Ashley Kendell is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Original photos by Dr. Julie Fleischman used by permission. Pubic symphyses are curated in the Hartnett-Fulginiti donated skeletal collection. Donation and research consent was provided by next of kin.]<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Degenerative changes in the skeleton typically begin after 18 years of age, with more prominent changes developing after an individual reaches middle adulthood (commonly defined as after 35 years of age in osteology). These changes are most easily seen around joint surfaces of the pelvis, the cranial vault, and the ribs. In this chapter, we focus on the pubic symphysis surfaces of the pelvis and the sternal ends of the ribs, which show metamorphic changes from young adulthood to older adulthood. The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1262\">pubic symphysis<\/a> <\/strong>is a joint that unites the left and right halves of the pelvis. The surface of the pubic symphysis changes during adulthood, beginning as a surface with pronounced ridges (called billowing) and flattening with a more distinct rim to the pubic symphysis as an individual ages. As with all metamorphic age changes, older adults tend to develop lipping around the joint surfaces as well as a breakdown of the joint surfaces. The most commonly used method for aging adult skeletons from the pubic symphysis is the Suchey-Brooks method (Brooks and Suchey 1990; Katz and Suchey 1986). This method divides the changes seen with the pubic symphysis into six phases based on macroscopic age-related changes to the surface. Figure 15.13 provides a visual of the degenerative changes that typically occur on the pubic symphysis.<\/p>\n<figure style=\"width: 403px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image12-3.png\" alt=\"Three sternal rib ends demonstrating progressive changes that occur with age.\" width=\"403\" height=\"220\" \/><figcaption class=\"wp-caption-text\">Figure 15.14: Examples of degenerative changes to the sternal rib end: (A) young adult; (B) middle adult; (C) old adult. Images derived from CSU, Chico HIL donated skeletal collection. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Examples of degenerative changes to the sternal rib end (Figure 15.15)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The sternal end of the ribs, the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1246\">anterior <\/a><\/strong> end of the rib that connects via cartilage to the sternum, is also used in age estimations of adults. This method, first developed by M. Y. \u0130\u015fcan and colleagues, considers both the change in shape of the sternal end as well as the quality of the bone (\u0130\u015fcan, Loth, and Wright 1984; \u0130\u015fcan, Loth, and Wright 1985). The sternal end first develops a billowing appearance in young adulthood. The bone typically develops a wider and deeper cupped end as an individual ages. Older adults tend to exhibit bony extensions of the sternal end rim as attaching cartilage ossifies. Figure 15.14 provides a visual of the degenerative changes that typically occur in sternal rib ends.<\/p>\n<h4 class=\"import-Normal\"><em>Estimating Stature<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Stature, or height, is one of the most prominently recorded components of the biological profile. Our height is recorded from infancy through adulthood. Doctor\u2019s appointments, driver's license applications, and sports rosters all typically involve a measure of stature for an individual. As such, it is also a component of the biological profile nearly every individual will have on record. Bioarchaeologists and forensic anthropologists use stature estimation methods to provide a range within which an individual\u2019s biological height would fall. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1248\">Biological height<\/a> <\/strong>is a person\u2019s true anatomical height. However, the range created through these estimations is often compared to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1264\">reported stature<\/a><\/strong>, which is typically self-reported and based on an approximation of an individual\u2019s true height (Ousley 1995).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In June 2015, two men were shot and killed in Granite Bay, California, in a double homicide. Investigators were able to locate surveillance camera footage from a gas station where the two victims were spotted in a car with another individual believed to be the perpetrator in the case. The suspect, sitting behind the victims in the car, hung his right arm out of the window as the car drove away. The search for the perpetrator was eventually narrowed down to two suspects. One suspect was 5\u2019 8\u201d while the other suspect was 6\u2019 4\u201d, representing almost a foot difference in height reported stature between the two. Forensic anthropologists were given the dimensions of the car (for proportionality of the arm) and were asked to calculate the stature of the suspect in the car from measurements of the suspect\u2019s forearm hanging from the window. Approximate lengths of the bones of the forearm were established from the video footage and used to create a predicted stature range. Stature estimations from skeletal remains typically look at the correlation between the measurements of any individual bone and the overall measurement of body height. In the case above, the length of the right forearm pointed to the taller of the two suspects who was subsequently arrested for the homicide.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Certain bones, such as the long bones of the leg, contribute more to our overall height than others and can be used with mathematical equations known as regression equations. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1252\">Regression methods <\/a> <\/strong>examine the relationship between variables such as height and bone length and use the correlation between the variables to create a prediction interval (or range) for estimated stature. This method for calculating stature is the most commonly used method (SWGANTH 2012). Figure 15.15 shows the measurement of the bicondylar length of the femur for stature estimations.<\/p>\n<figure style=\"width: 584px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image2-3.png\" alt=\"A femur is measured using a wooden osteometric board.\" width=\"584\" height=\"389\" \/><figcaption class=\"wp-caption-text\">Figure 15.15: Image of measurement of the bicondylar length of the femur, often used in the estimation of living stature. Image derived from CSU, Chico HIL donated skeletal collection. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Measurement of the bicondylar length of the femur<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Identification Using Individualizing Characteristics<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">One of the most frequently requested analyses within the forensic anthropology laboratory is assistance with the identification of unidentified remains. While all components of a biological profile, as discussed above, can assist law enforcement officers and medical examiners to narrow down the list of potential identifications, a biological profile will not lead to a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1254\">positive identification<\/a><\/strong>. The term <em>positive identification<\/em> refers to a scientifically validated method of identifying previously unidentified remains. Presumptive identifications, however, are not scientifically validated; rather, they are based on circumstances or scene context. For example, if a decedent is found in a locked home with no evidence of forced entry but the body is no longer visually identifiable, it may be presumed that the remains belong to the homeowner. Hence, a presumptive identification.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The medicolegal system ultimately requires that a positive identification be made in such circumstances, and a presumptive identification is often a good way to narrow down the pool of possibilities. Biological profile information also assists with making a presumptive identification based on an individual\u2019s phenotype in life (e.g., what they looked like). As an example, a forensic anthropologist may establish the following components of a biological profile: white male, between the ages of 35 and 50, approximately 5\u2019 7\u201d to 5\u2019 11.\u201d While this seems like a rather specific description of an individual, you can imagine that this description fits dozens, if not hundreds, of people in an urban area. Therefore, law enforcement can use the biological profile information to narrow their pool of possible identifications to include only white males who fit the age and height outlined above. Once a possible match is found, the decedent can be identified using a method of positive identification.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Positive identifications are based on what we refer to as individualizing traits or characteristics, which are traits that are unique at the individual level. For example, brown hair is not an individualizing trait as brown is the most common hair color in the U.S. But, a specific pattern of dental restorations or surgical implants can be individualizing, because it is unlikely that you will have an exact match on either of these traits when comparing two individuals.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">A number of positive methods are available to forensic anthropologists, and for the remainder of this section we will discuss the following methods: comparative medical and dental radiography and identification of surgical implants.<\/p>\n<figure style=\"width: 165px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image17-3.png\" alt=\"Radiograph of skull with frontal sinuses visible.\" width=\"165\" height=\"182\" \/><figcaption class=\"wp-caption-text\">Figure 15.16: Example of the unique shape of the frontal sinus. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Frontal_bone_sinuses.jpg\">Frontal bone sinuses<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Alex_Khimich\">Alex Khimich<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Comparative medical and dental radiography is used to find consistency of traits when comparing antemortem records (medical and dental records taken during life) with images taken postmortem (after death). Comparative medical radiography focuses primarily on features associated with the skeletal system, including trabecular pattern (internal structure of bone that is honeycomb in appearance), bone shape or cortical density (compact outer layer of bone), and evidence of past trauma, skeletal pathology, or skeletal anomalies. Other individualizing traits include the shape of various bones or their features, such as the frontal sinuses (Figure 15.16).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Comparative dental radiography focuses on the number, shape, location, and orientation of dentition and dental restorations in antemortem and postmortem images. While there is not a minimum number of matching traits that need to be identified for an identification to be made, the antemortem and postmortem records should have enough skeletal or dental consistencies to conclude that the records did in fact come from the same individual (SWGANTH 2010a). Consideration should also be given to population-level frequencies of specific skeletal and dental traits. If a trait is particularly common within a given population, it may not be a good trait to utilize for positive identification.<\/p>\n<figure style=\"width: 354px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image16-3.png\" alt=\"A scapula and humerus with a metal shoulder replacement.\" width=\"354\" height=\"231\" \/><figcaption class=\"wp-caption-text\">Figure 15.17: Image of joint replacement in the right shoulder. Credit: <a href=\"https:\/\/naturalhistory.si.edu\/education\/teaching-resources\/written-bone\/skeleton-keys\/todays-bones\">Shoulder replacement<\/a> by <a href=\"https:\/\/www.si.edu\/\">Smithsonian<\/a> [exhibit: Written in Bone, Today\u2019s Bones] <a href=\"https:\/\/www.si.edu\/termsofuse\">is used for educational and non-commercial purposes as outlined by the Smithsonian.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Surgical implants or devices can also be used for identification purposes (Figure 15.17). These implements are sometimes recovered with human remains. One of the ways forensic anthropologists can use surgical implants to assist in decedent identification is by providing a thorough analysis of the implant and noting any identifying information such as serial numbers, manufacturer symbols, and so forth. This information can then sometimes be tracked directly to the manufacturer or the place of surgical intervention, which may be used to identify unknown remains (SWGANTH 2010a).<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Trans Doe Task Force<\/h2>\n<p class=\"import-Normal\">The Trans Doe Task Force (TDTF) is a Trans-led nonprofit organization that investigates cases involving LGBTQ+ missing and murdered persons. The organization specifically focuses on transgender and gender-variant cases, providing connections between law enforcement agencies, medical examiner offices, forensic anthropologists, and forensic genetic genealogists to increase the chances of identification. Additionally, the TDTF curates a data repository of missing, murdered, and unclaimed LGBTQ+ individuals, and they continuously try innovative approaches to identify these individuals, whose lived gender identity may not match their biological sex.<\/p>\n<p class=\"import-Normal\">For more information visit <a href=\"https:\/\/transdoetaskforce.org\/\">transdoetaskforce.org<\/a><\/p>\n<\/div>\n<h3 class=\"import-Normal\"><strong>Trauma Analysis<\/strong><\/h3>\n<h4 class=\"import-Normal\"><em>Types of Trauma<\/em><strong><br \/>\n<\/strong><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Within the field of anthropology, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1256\">trauma<\/a> <\/strong>is defined as an injury to living tissue caused by an extrinsic force or mechanism (Lovell 1997:139). Forensic anthropologists can assist a forensic pathologist by providing an interpretation of the course of events that led to skeletal trauma. Typically, traumatic injury to bone is classified into one of four categories, defined by the trauma mechanism. A trauma mechanism refers to the force that produced the skeletal modification and can be classified as (1) sharp force, (2) blunt force, (3) projectile, or (4) thermal (burning). Each type of trauma, and the characteristic pattern(s) associated with that particular categorization, will be discussed below.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">First, let\u2019s consider s<em>harp-force trauma<\/em>, which is caused by a tool that is edged, pointed, or beveled\u2014for example, a knife, saw, or machete (SWGANTH 2011). The patterns of injury resulting from sharp-force trauma include linear incisions created by a sharp, straight edge; punctures; and chop marks (Figure 15.18; SWGANTH 2011). When observed under a microscope, an anthropologist can often determine what kind of tool created the bone trauma. For example, a power saw cut will be discernible from a manual saw cut.<\/p>\n<figure style=\"width: 602px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image21-1.png\" alt=\"Anterior image of a skull with multiple traumatic injuries to forehead.\" width=\"602\" height=\"457\" \/><figcaption class=\"wp-caption-text\">Figure 15.18: Example of sharp-force trauma (sword wound) to the frontal bone. The skull appears sliced with thin lines in two places across the top of the skull. Credit: <a href=\"https:\/\/openverse.org\/image\/909d1b77-ad5f-4cda-be44-6d9b5fbf14b9\/\">Female skull injured by a medieval sword<\/a> by <a href=\"https:\/\/sketchfab.com\/provinciaal_depot_noordholland\">Provinciaal depot voor archeologie Noord-Holland<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY 4.0 License<\/a>. The original image is a 3D model that can be manipulated on the <a href=\"https:\/\/wordpress.org\/openverse\/image\/909d1b77-ad5f-4cda-be44-6d9b5fbf14b9\/\">openverse website<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Second, <em>blunt-force trauma<\/em> is defined as \u201ca relatively low-velocity impact over a relatively large surface area\u201d (Galloway 1999, 5). Blunt-force injuries can result from impacts from clubs, sticks, fists, and so forth. Blunt-force impacts typically leave an injury at the point of impact but can also lead to bending and deformation in other regions of the bone. Depressions, fractures, and deformation at and around the site of impact are all characteristics of blunt-force trauma (Figure 15.19). As with sharp-force trauma, an anthropologist attempts to interpret blunt-force injuries, providing information pertaining to the type of tool used, the direction of impact, the sequence of impacts, if more than one, and the amount of force applied.<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 578px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image30.png\" alt=\"Cranium with two blunt force impacts from a hammer.\" width=\"578\" height=\"803\" \/><figcaption class=\"wp-caption-text\">Figure 15.19: Example of multiple blunt force impacts to the left parietal and frontal bones. There is one hole in the skull with fractured bone around the edges. There are also multiple spots across the back of the skull with depressions of various sizes. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Skull_hammer_trauma.jpg\">Skull hammer trauma<\/a> by <a href=\"https:\/\/www.nih.gov\/\">the National Institutes of Health<\/a>, Health &amp; Human Services, is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>. [Exhibit: Visible Proofs: Forensic Views of the Body, U.S. National Library of Medicine, 19th Century Collection, National Museum of Health and Medicine, Armed Forces Institute of Pathology, Washington, D.C.]<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Third, <em>projectile trauma<\/em> refers to high-velocity trauma, typically affecting a small surface area (Galloway 1999, 6). Projectile trauma results from fast-moving objects such as bullets or shrapnel. It is typically characterized by penetrating defects or embedded materials (Figure 15.20). When interpreting injuries resulting from projectile trauma, an anthropologist can often offer information pertaining to the type of weapon used (e.g., rifle vs. handgun), relative size of the bullet (but not the caliber of the bullet), the direction the projectile was traveling, and the sequence of injuries if there are multiple present.<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 462px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image5-3.png\" alt=\"Anterior and posterior views of a skull with a gunshot wound.\" width=\"462\" height=\"291\" \/><figcaption class=\"wp-caption-text\">Figure 15.20: Example of projectile trauma with an entrance wound to the frontal bone and exit wound visible on the occipital. A small circular hole is visible in the front of the skull with cracks radiating out from the point of impact. There is a larger hole visible in the back of the skull that is irregular yet circular in shape. Credit: <a href=\"https:\/\/naturalhistory.si.edu\/education\/teaching-resources\/written-bone\/skeleton-keys\/how-bone-biographies-get-written\">Trauma: Gunshot Wounds<\/a> by <a href=\"https:\/\/www.si.edu\/\">Smithsonian<\/a> [exhibit: Written in Bone, How Bone Biographies Get Written] <a href=\"https:\/\/www.si.edu\/termsofuse\">is used for educational and non-commercial purposes as outlined by the Smithsonian.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Finally, <em>thermal trauma<\/em> is a bone alteration that results from bone exposure to extreme heat. Thermal trauma can result in cases of house or car fires, intentional disposal of a body in cases of homicidal violence, plane crashes, and so on. Thermal trauma is most often characterized by color changes to bone, ranging from yellow to black (charred) or white (calcined). Other bone alterations characteristic of thermal trauma include delamination (flaking or layering due to bone failure), shrinkage, fractures, and heat-specific burn patterning. When interpreting injuries resulting from thermal damage, an anthropologist can differentiate between thermal fractures and fractures that occurred before heat exposure, thereby contributing to the interpretation of burn patterning (e.g., was the individual bound or in a flexed position prior to the fire?).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">While there are characteristic patterns associated with the four categories of bone trauma, it is also important to note that these bone alterations do not always occur independently of different trauma types. An individual\u2019s skeleton may present with multiple different types of trauma, such as a projectile wound and thermal trauma. Therefore, it is important that the anthropologist recognize the different types of trauma and interpret them appropriately.<\/p>\n<h3 class=\"import-Normal\"><strong>Timing of Injury<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Another important component of any anthropological trauma analysis is the determination of the timing of injury (e.g., when did the injury occur). Timing of injury is traditionally split into one of three categories: <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1260\">antemortem<\/a> <\/strong>(before death), <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1258\">perimortem<\/a> <\/strong>(at or around the time of death), and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1266\">postmortem <\/a><\/strong>(after death). This classification system differs slightly from the classification system used by the pathologist because it specifically references the qualities of bone tissue and bone response to external forces. Therefore, the perimortem interval (at or around the time of death) means that the bone is still fresh and has what is referred to as a green bone response, which can extend past death by several weeks or even months. For example, in cold or freezing temperatures a body can be preserved for extended periods of time, increasing the perimortem interval, while in desert climates decomposition is accelerated, thereby significantly decreasing the postmortem interval (Galloway 1999, 12). Antemortem injuries (occurring well before death and not related to the death incident) are typically characterized by some level of healing, in the form of a fracture callus or unification of fracture margins. Finally, postmortem injuries (occurring after death, while bone is no longer fresh) are characterized by jagged fracture margins, resulting from a loss of moisture content during the decomposition process (Galloway 1999, 16). In general, all bone traumas should be classified according to the timing of injury, if possible. This information will help the medical examiner or pathologist better understand the circumstances surrounding the decedent\u2019s death, as well as events occurring during life and after the final disposition of the body.<\/p>\n<h3 class=\"import-Normal\"><strong>The Role of the Forensic Anthropologist in Trauma Analysis<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Within the medicolegal system, forensic anthropologists are often called upon by the medical examiner, forensic pathologist, or coroner to assist with an interpretation of trauma. The forensic anthropologist\u2019s main focus in any trauma analysis is the underlying skeletal system\u2014as well as, sometimes, cartilage. Analysis and interpretation of soft tissue injuries fall within the purview of the medical examiner or pathologist. It is also important to note that the main role of the forensic anthropologist is to provide information pertaining to skeletal injury to assist the medical examiner\/pathologist in their final interpretation of injury. Forensic anthropologists do not hypothesize as to the cause of death of an individual. Instead, a forensic anthropologist\u2019s report should include a description of the injury (e.g., trauma mechanism, number of injuries, location, timing of injury); documentation of the injury, which may be utilized in court testimony (e.g., photographs, radiographs, measurements); and, if applicable, a statement as to the condition of the body and state of decomposition, which may be useful for understanding the depositional context (e.g., how long has the body been exposed to the elements; was it moved or in its original location; are any of the alterations to bone due to environmental or faunal exposure instead of intentional human modification).<\/p>\n<h2 class=\"import-Normal\">Taphonomy<\/h2>\n<h2 class=\"import-Normal\"><strong>What Happened to the Remains After Death?<\/strong><\/h2>\n<p class=\"import-Normal\">The majority of the skeletal analysis process revolves around the identity of the deceased individual. However, there is one last, very important question that forensic anthropologists should ask: What happened to the remains after death? Generally speaking, processes that alter the bone after death are referred to as taphonomic changes (refer to Chapter 7 for a discussion regarding taphonomy and the fossil record).<\/p>\n<p class=\"import-Normal\">The term <em>taphonomy<\/em> was originally used to refer to the processes through which organic remains mineralize, also known as fossilization. Within the context of biological anthropology, the term <em>taphonomy<\/em> is better defined as the study of what happens to human remains after death (Komar and Buikstra 2008). Initial factors affecting a body after death include processes such as decomposition and scavenging by animals. However, taphonomic processes encompass much more than the initial period after death. For example, plant root growth can leach minerals from bone, leaving a distinctive mark. Sunlight can bleach human remains, leaving exposed areas whiter than those that remained buried. Water can wear the surface of the bone until it becomes smooth.<\/p>\n<p class=\"import-Normal\">Some taphonomic processes can help a forensic anthropologist estimate the relative amount of time that human remains have been exposed to the elements. For example, root growth through a bone would certainly indicate a body was buried for more than a few days. Forensic anthropologists must be very careful when attempting to estimate time since death based on taphonomic processes because environmental conditions can greatly influence the rate at which taphonomic processes progress. For example, in cold environments, tissue may decay slower than in warm, moist environments.<\/p>\n<p class=\"import-Normal\">Forensic anthropologists must contend with taphonomic processes that affect the preservation of bones. For example, high acidity in the soil can break down human bone to the point of crumbling. In addition, when noting trauma, they must be very careful not to confuse postmortem (after death) bone damage with trauma.<\/p>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 470.25pt\">\n<caption>Figure 15.21: Table showing taphonomic processes that affect the preservation of bones. A. Rodent gnawing. B. Carnivore damage. C. Burned bone. D. Root etching. E. Weathering. F. Cut marks. Credit: A. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Rodent gnawing (Figure 15.26)<\/a>, B. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Carnivore damage (Figure 15.27)<\/a>, C. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Burned bone (Figure 15.28)<\/a>, D. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Root etching (Figure 15.29)<\/a>, E. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Weathering (Figure 15.30)<\/a>, and F. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Cut marks (Figure 15.30)<\/a>, all original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone are under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-NC 4.0 License<\/a>.<\/caption>\n<thead>\n<tr style=\"height: 52.5pt\">\n<td class=\"Table1-C\" style=\"padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Taphonomic Process<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 1pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Definition<\/p>\n<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table1-R\" style=\"height: 190.5pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Rodent Gnawing<\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: center\"><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image19-2.png\" alt=\"Parallel tooth marks etched by a rodent\u2019s front teeth visible on the end of an animal bone.\" width=\"564\" height=\"422\" \/><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">When rodents, such as rats and mice, chew on bone, they leave sets of parallel grooves. The shallow grooves are etched by the rodent\u2019s incisors.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 166.75pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Carnivore Damage<\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: center\"><strong><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image23-4.png\" alt=\"Pit marks from the canines of a carnivore visible on the surface of an animal bone.\" width=\"410\" height=\"272\" \/><\/strong><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Carnivores may leave destructive dental marks on bone. The tooth marks may be visible as pit marks or punctures from the canines, as well as extensive gnawing or chewing of the ends of the bones to retrieve marrow.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 177pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Burned Bone<\/strong><\/p>\n<p class=\"import-Normal\"><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image20-5.png\" alt=\"Burned animal bone fragments pictured at different stages of thermal damage.\" width=\"512\" height=\"342\" \/><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Fire causes observable damage to bone. Temperature and the amount of time bone is heated affect the appearance of the bone. Very high temperatures can crack bone and result in white coloration. Color gradients are visible in between high and lower temperatures, with lower temperatures resulting in black coloration from charring. Cracking can also reveal information about the directionality of the burn.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 169.75pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Root Etching<\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: center\"><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image15-4.png\" alt=\"Animal bone with prominent, discolored grooves where roots leached nutrients from bone\u2019s surface.\" width=\"512\" height=\"342\" \/><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Plant roots can etch the outer surface of bone, leaving grooves where the roots attached as they leached nutrients. During this process, the plant\u2019s roots secrete acid that breaks down the surface of the bone.<\/p>\n<p class=\"import-Normal\">\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 170.5pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Weathering<\/strong><\/p>\n<p class=\"import-Normal\"><strong><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image9.png\" alt=\"Cracking and exfoliation of the surface of an animal bone. \" width=\"512\" height=\"342\" \/><\/strong><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Many different environmental conditions affect bone. River transport can smooth the surface of the bone due to water abrasion. Sunlight can bleach the exposed surface of bone. Dry and wet environments or the mixture of both types of environments can cause cracking and exfoliation of the surface. Burial in different types of soil can cause discoloration, and exposure can cause degreasing.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 169.75pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Cut Marks<\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: left\"><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image8-2.png\" alt=\"Thin vertical lines and cuts are visible along the bone.\" width=\"512\" height=\"342\" \/><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Humans may alter bone by cutting, scraping, or sawing it directly or in the process of removing tissue. The groove pattern\u2014that is, the depth and width of the cuts\u2014can help identify the tool used in the cutting process.<\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<h2>Dig Deeper: Modern Forensic Technologies<\/h2>\n<p>In recent years, the forensics community has greatly benefited from the introduction of new technologies, helping strengthen the precision and speed of discoveries and advancements in the field. With recent developments in forensic anthropology, such as 3D scanning technologies, virtual reconstruction, and AI-assisted DNA analysis being integrated into traditional methods, there have been notable changes in how experts investigate human remains.<\/p>\n<p><strong>Artificial intelligence<\/strong><\/p>\n<p>In recent years, Artificial intelligence (AI) has shown itself to be a valuable tool within forensic anthropology. Aiding forensic experts and toxicologists with complex tasks, the limitations of traditional autopsies can be addressed with the help of AI. By automating and enhancing key investigative processes such as searching for microscopic changes in the human body to determine the cause of death or a person\u2019s life conditions, AI has the potential to enhance the efficiency of forensic processes significantly. It facilitates the detection of microscopic bodily changes to determine the cause of death or living conditions, compares evidence against databases for weapon identification and blood spatter analysis, and reduces manual workload. AI also enables the electronic storage of biometric data\u2013such as facial features, retinal patterns, and fingerprints\u2013for more accurate identity verification. Additionally, AI-powered microscopy enhances the detection of biological traces on complex surfaces, while blood biomarker analysis allows for more precise estimations of time of death (Wankhade et al., 2022).<\/p>\n<p>While AI holds great promise for the future of forensic medicine, a significant challenge remains: sourcing high-quality data to train the algorithms effectively. One of the more recent AI technologies making waves in the forensic anthropology sector is a new automated AI algorithm called the Convolutional Neural Network (CNN). As described by researchers in Switzerland\u2019s national medical journal Healthcare, CNN is a Deep Learning algorithm that allows for the detection of microscopic skull damage from CT scans or soft-tissue predictions of a face based on the skull information provided (Thurzo et al., 2021). While there are many advantages to using the CNN, the algorithm can be subject to biases in the same way human forensic experts can, as its assessment and pattern recognition of skulls and skeletons depend on the source data initially used for its AI training (2021).<\/p>\n<p><strong>3D Modeling<\/strong><\/p>\n<p>Identifying complex trauma to bones\u2013such as distinguishing heat fractures following blunt force trauma\u2013remains a significant challenge in forensic anthropology. This is particularly true for irregular skeletal structures like the pelvis, where overlapping trauma types can be difficult to differentiate, leading to these bones often being understudied. A 2024 study done by researchers from the University of Alberta in collaboration with the Michigan State Police explores the use of 3D laser scans and modelling technology to provide a highly detailed analysis of irregular bones with trauma. The study aimed to better distinguish peri-mortem trauma (trauma occurring around the time of death) from post-mortem heat alterations and improve the forensic analysis accuracy of such cases (Friedlander et al., 2024). The use of 3D laser scans and modelling technology provides very clear, detailed, and colored scans of bones, showing distinctions between the characteristics of the fractures. Blunt force and sharp force trauma produce a colour gradient on the 3D model that is more gradual and irregular, while heat fractures are more neat and characterized by little colour variation on the 3D models (2024). Other conclusions were also drawn from the study, such as the differences in trauma on fresh bones and bones that have been exposed to the elements for longer. An example of this is the interstitial fluid and collagen fibrils in fresh bones absorbing force, causing more long and jagged fracture lines, as opposed to a brittle fracture that older bones may exhibit (2024).<\/p>\n<p>Overall, the integration of 3D modeling technology offers a reproducible and highly detailed approach for analyzing trauma in anatomically complex and historically understudied skeletal regions. The practicality of this advancement is further emphasized by the researchers, who note that \u201cin many instances, scanned 3D models can be 3D printed for handheld representation of the model without damaging or overhandling the remains\u201d (2024, p. 2). By enhancing the ability to differentiate between various types of trauma and allowing for more convenient and risk-averse methods of research, this technology significantly improves the accuracy and reliability of forensic interpretations.<\/p>\n<h2>Ethics and Human Rights<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Working with human remains requires a great deal of consideration and respect for the dead. Forensic anthropologists have to think about the ethics of our use of human remains for scientific purposes. How do we conduct casework in the most respectable manner possible? While there are a wide range of ethical considerations to consider when contemplating a career in forensic anthropology, this chapter will focus on two major categories: working with human remains and acting as an expert within the medicolegal system.<\/p>\n<h3 class=\"import-Normal\"><strong>Working with Human Remains<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropologists work with human remains in a number of contexts, including casework, excavation, research, and teaching. When working with human remains, it is always important to use proper handling techniques. To prevent damage to skeletal remains, bones should be handled over padded surfaces. Skulls should never be picked up by placing fingers in the eye orbits, foramen magnum (hole at the base of the skull for entry of the spinal cord), or through the zygomatic arches (cheekbones). Human remains, whether related to casework, fieldwork, donated skeletal collections, or research, were once living human beings. It is important to always bear in mind that work with remains should be ingrained with respect for the individual and their relatives. In addition to fieldwork, casework, and teaching, anthropologists are often invited to work with remains that come from a bioarchaeological context or from a human rights violation. While this discussion of ethics is not comprehensive, two case examples will be provided below in which an anthropologist must consider the ethical standards outlined above.<\/p>\n<h3 class=\"import-Normal\"><strong>Modern Human Rights Violations<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropologists may also be called to participate in criminal investigations involving human rights violations. Anthropological investigations may include assistance with identifications, determination of the number of victims, and trauma analyses. In this role, forensic anthropologists play an integral part in promoting human rights, preventing future human rights violations, and providing the evidence necessary to prosecute those responsible for past events. A few ethical considerations for the forensic anthropologist involved in human rights violations include the use of appropriate standards of identification, presenting reliable and unbiased testimony, and maintaining preservation of evidence. For a more comprehensive history of forensic anthropological contributions to human rights violations investigations, see Ubelaker 2018.<\/p>\n<h3 class=\"import-Normal\"><strong>Acting as an Expert in the Medicolegal System<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In addition to the ethical considerations involved in working with human skeletal remains, forensic anthropologists must abide by ethical standards when they act as experts within the medicolegal system. The role of the forensic anthropologist within the medicolegal system is primarily to provide information to the medical examiner or coroner that will aid in the identification process or determination of cause and manner of death. Forensic anthropologists also may be called to testify in a court of law. In this capacity, forensic anthropologists should always abide by a series of ethical guidelines that pertain to their interpretation, presentation, and preservation of evidence used in criminal investigations. First and foremost, practitioners should never misrepresent their training or education. When appropriate, outside opinions and assistance in casework should be requested (e.g., consulting a radiologist for radiological examinations or odontologist for dental exams). The best interest of the decedent should always take precedence. All casework should be conducted in an unbiased way, and financial compensation should never be accepted as it can act as an incentive to take a biased stance regarding casework. All anthropological findings should be kept confidential, and release of information is best done by the medical examiner or coroner. Finally, while upholding personal ethical standards, forensic anthropologists are also expected to report any perceived ethical violations committed by their peers.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ethical standards for the field of forensic anthropology are outlined by the Organization of Scientific Area Committees (OSAC) for Forensic Science, administered by the National Institute of Standards and Technology (NIST). OSAC and NIST recently began an initiative to develop standards that would strengthen the practice of forensic science both in the United States and internationally. OSAC\u2019s main objective is to \u201cstrengthen the nation\u2019s use of forensic science by facilitating the development of technically sound forensic science standards and by promoting the adoption of those standards by the forensic science community\u201d (NIST n.d.). Additionally, OSAC promotes the establishment of best practices and other guidelines to ensure that forensic science findings and their presentation are reliable and reproducible (NIST 2023).<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Native American Graves Protection and Repatriation Act (NAGPRA)<\/h2>\n<p class=\"import-Normal\">There is a long history in the <span style=\"background-color: #00ffff\">United States<\/span> of systematic disenfranchisement of Native American people, including lack of respect for tribal sovereignty. This includes the egregious treatment of Native American human remains. Over several centuries, thousands of Native American remains were removed from tribal lands and held at institutions in the United States, such as museums and universities.<\/p>\n<p class=\"import-Normal\">In 1990, a landmark human rights federal law, the Native American Graves Protection and Repatriation Act (NAGPRA), spurred change in the professional standards and practice of biological anthropology and archaeology. NAGPRA established a legal avenue to provide protection for and repatriation of Native American remains, cultural items, and sacred objects removed from Federal or tribal lands to Native American lineal descendants, Indian tribes, and Native Hawaiian organizations. Human remains and associated artifacts, curated in museum collections and federally funded institutions, are subject to three primary provisions outlined by the NAGPRA statute: (1) protection for Native graves on federal and private land; (2) recognition of tribal authority on such lands; and (3) the requirement that all Native skeletal remains and associated artifacts be inventoried and culturally affiliated groups be consulted concerning decisions related to ownership and final disposition (Rose, Green, and Green 1996). NAGPRA legislation was enacted to ensure ethical consideration and treatment of Native remains and to improve dialogue between scientists and Native groups.<\/p>\n<ul>\n<li>For more information about NAGPRA, visit the <a href=\"https:\/\/www.usbr.gov\/nagpra\/\" target=\"_blank\" rel=\"noopener\">Bureau of Reclamation NAGPRA website<\/a><\/li>\n<li>To read the text of the law, visit the <a href=\"https:\/\/www.congress.gov\/bill\/101st-congress\/house-bill\/5237\">US Congress NAGPRA law website<\/a>.<\/li>\n<li>For further discussion of NAGPRA history, please see <a href=\"https:\/\/textbooks.whatcom.edu\/tracesarchaeology\/\" target=\"_blank\" rel=\"noopener\"><em>TRACES: <\/em><em>An Open Invitation to <\/em><em>Archaeology <\/em>open textbook website<\/a><em><br \/>\n<\/em><\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Becoming a Forensic Anthropologist<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"background-color: #ff99cc\">What does it take to be a forensic anthropologist? Forensic anthropologists are first and foremost anthropologists. While many forensic anthropologists have an undergraduate degree in anthropology, they may also major in biology, criminal justice, pre-law, pre-med, and many other related fields. Practicing forensic anthropologists typically have an advanced degree, either a Master\u2019s or Doctoral degree in Anthropology. Additional training and experience in archaeology, the medico-legal system, rules of evidence, and expert witness testimony are also common. Practicing forensic anthropologists are also encouraged to be board-certified through the American Board of Forensic Anthropology (ABFA). Learn more about the field and educational opportunities on the ABFA website: <a class=\"rId111\" style=\"background-color: #ff99cc\" href=\"https:\/\/www.theabfa.org\/coursework\">https:\/\/www.theabfa.org\/coursework<\/a>.<\/span><\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Review Questions<\/h2>\n<ul>\n<li>What is forensic anthropology? What are the seven primary steps involved in a skeletal analysis?<\/li>\n<li>What are the major components of a biological profile? Why are forensic anthropologists often-tasked with creating biological profiles for unknown individuals?<\/li>\n<li>What are the four major types of skeletal trauma?<\/li>\n<li>What is taphonomy, and why is an understanding of taphonomy often critical in forensic anthropology analyses?<\/li>\n<li>What are some of the ethical considerations faced by forensic anthropologists?<\/li>\n<\/ul>\n<\/div>\n<h2>About the Authors<\/h2>\n<p><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image24-4.jpg\" alt=\"A woman with straight blonde hair smiles at the camera. \" width=\"191\" height=\"254\" \/><\/p>\n<h3 class=\"import-Normal\"><strong>Ashley Kendell, Ph.D.<\/strong><\/h3>\n<p class=\"import-Normal\">California State University, Chico, <a class=\"rId113\" href=\"mailto:akendell@csuchico.edu\">akendell@csuchico.edu<\/a><\/p>\n<p class=\"import-Normal\">Dr. Ashley Kendell is currently an associate professor and forensic anthropologist at Chico State. Prior to beginning her position at Chico State, she was a visiting professor at the University of Montana and the forensic anthropologist for the state of Montana. Dr. Kendell obtained her doctorate from Michigan State University, and her research interests include skeletal trauma analysis and digitization and curation methods for digital osteological data. She is also a Registry Diplomate of the American Board of Medicolegal Death Investigators. Throughout her doctoral program, she worked as a medicolegal death investigator for the greater Lansing, Michigan, area and was involved in the investigation of over 200 forensic cases.<\/p>\n<p class=\"import-Normal\"><strong><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image4.jpg\" alt=\"A woman with straight brown hair pulled back smiles at the camera. \" width=\"194\" height=\"258\" \/><\/strong><\/p>\n<h3 class=\"import-Normal\"><strong>Alex Perrone, M.A., M.S.N, R.N., P.H.N.<\/strong><\/h3>\n<p class=\"import-Normal\">Butte Community College, <a class=\"rId115\" href=\"mailto:perroneal@butte.edu\">perroneal@butte.edu<\/a><\/p>\n<p class=\"import-Normal\">Alex Perrone is a lecturer in anthropology at Butte Community College. She is also a Registered Nurse and a certified Public Health Nurse. She is a former Supervisor of the Human Identification Laboratory in the Department of Anthropology at California State University, Chico. Her research interests include bioarchaeology, paleopathology, forensic anthropology, skeletal biology, California prehistory, and public health. She has worked on bioarchaeological and archaeological projects in Antigua, California, Hawaii, Greece, and the UK, and was an archaeological technician for the USDA Forest Service. She assisted with training courses for local and federal law enforcement agencies and assisted law enforcement agencies with the recovery and analysis of human remains.<\/p>\n<p class=\"import-Normal\" data-wp-editing=\"1\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image1-1.jpg\" alt=\"A woman with curly brown, shoulder-length hair smiles at the camera.\" width=\"190\" height=\"253\" \/><\/p>\n<h3 class=\"import-Normal\"><strong>Colleen Milligan, Ph.D.<\/strong><\/h3>\n<p class=\"import-Normal\">California State University, Chico, <a class=\"rId117\" href=\"mailto:cfmilligan@csuchico.edu\">cfmilligan@csuchico.edu<\/a><\/p>\n<p class=\"import-Normal\">Dr. Colleen Milligan is a biological and forensic anthropologist with research interests in bioarchaeology, skeletal biology, and forensic anthropology. She has been a Fellow with the Department of Homeland Security and has assisted in forensic anthropology casework and recoveries in the State of Michigan and California. She has also assisted in community outreach programs in forensic anthropology and forensic science, as well as recovery training courses for local, state, and federal law enforcement officers. She is a certified instructor through Peace Officers Standards and Training (POST). Dr. Milligan serves as the current co-director of the Chico State Human Identification Laboratory.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration<\/h2>\n<p><a href=\"https:\/\/www.theabfa.org\/coursework\" target=\"_blank\" rel=\"noopener\">The American Board of Forensic Anthropology (ABFA)<\/a><\/p>\n<p><a href=\"https:\/\/www.aafs.org\/\" target=\"_blank\" rel=\"noopener\">The American Academy of Forensic Sciences (AAFS)<\/a><\/p>\n<p><a href=\"https:\/\/www.nist.gov\/organization-scientific-area-committees-forensic-science\" target=\"_blank\" rel=\"noopener\">The Organization of Scientific Area Committees for Forensic Science (OSAC)<\/a><\/p>\n<p><a href=\"https:\/\/textbooks.whatcom.edu\/tracesarchaeology\/\" target=\"_blank\" rel=\"noopener\">TRACES Bioarchaeology<\/a><\/p>\n<p><a href=\"https:\/\/transdoetaskforce.org\/\" target=\"_blank\" rel=\"noopener\">Trans Doe Task Force<\/a><\/p>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Adams, Bradley J., and Lyle W. Konigsberg, eds. 2008. <em>Recovery, Analysis, and Identification of Commingled Remains<\/em>. Totowa, NJ: Humana Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Beatrice, Jared S., and Angela Soler. 2016. \u201cSkeletal Indicators of Stress: A Component of the Biocultural Profile of Undocumented Migrants in Southern Arizona.\u201d <em>Journal of Forensic Sciences <\/em>61 (5): 1164\u20131172.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Berg, Gregory E. 2017. \u201cSex Estimation of Unknown Human Skeletal Remains.\u201d In <em>Forensic Anthropology: A Comprehensive Introduction, Second Edition<\/em>, edited by Natalie R. Langley and MariaTeresa A. Tersigni-Tarrant, 143\u2013159. Boca Raton, FL: CRC Press.<\/p>\n<p class=\"import-Normal\">Bethard, Jonathan D., and Elizabeth A. DiGangi. 2020. \u201cLetter to the Editor\u2014Moving Beyond a Lost Cause: Forensic Anthropology and Ancestry Estimates in the United States.\u201d <em>Journal of Forensic Sciences<\/em> 65 (5): 1791\u20131792.<\/p>\n<p class=\"import-Normal\">Birkby, Walter H., Todd W. Fenton, and Bruce E. Anderson. 2008. \u201cIdentifying Southwest Hispanics Using Nonmetric Traits and the Cultural Profile.\u201d <em>Journal of Forensic Sciences <\/em>53 (1): 29\u201333.<\/p>\n<p class=\"import-Normal\">Blatt, Samantha, Amy Michael, and Lisa Bright. Forthcoming. \u201cBioarchaeology: Interpreting Human Behavior from Skeletal Remains.\u201d In <em>TRACES: <\/em><em>An Open Invitation to <\/em><em>Archaeology<\/em>. https:\/\/textbooks.whatcom.edu\/tracesarchaeology\/.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Brooks, S., and J. M. Suchey. 1990. \u201cSkeletal Age Determination Based on the Os Pubis: A Comparison of the Acs\u00e1di-Nemesk\u00e9ri and Suchey-Brooks Methods.\u201d <em>Human Evolution <\/em>5 (3): 227\u2013238.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Buchanan, Shelby. 2014. \u201cBone Modification in Male to Female Transgender Surgeries: Considerations for the Forensic Anthropologist.\u201d MA thesis, Department of Geography and Anthropology, Louisiana State University, Baton Rouge.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Cunningham, Craig, Louise Scheuer, and Sue Black. 2016. <em>Developmental Juvenile Osteology, Second Edition<\/em>. London: Elsevier Academic Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Galloway, Alison, ed. 1999. <em>Broken Bones: Anthropological Analysis of Blunt Force Trauma<\/em>. Springfield, IL: Charles C. Thomas Publisher, LTD.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Hefner, Joseph T., and Kandus C. Linde. 2018. <em>Atlas of Human Cranial <\/em><em>Macromorphoscopic<\/em><em> Traits<\/em>. San Diego: Academic Press.<\/p>\n<p class=\"import-Normal\">\u0130\u015fcan, M. Y., S. R. Loth, and R. K. Wright. 1984. \u201cAge Estimation from the Rib by Phase Analysis: White Males.\u201d <em>Journal of Forensic Sciences <\/em>29 (4): 1094\u20131104.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">\u0130\u015fcan, M. Y., S. R. Loth, and R. K. Wright. 1985. \u201cAge Estimation from the Rib by Phase Analysis: White Females.\u201d <em>Journal of Forensic Sciences <\/em>30 (3): 853\u2013863.Katz, Darryl, and Judy Myers Suchey. 1986. \u201cAge Determination of the Male Os Pubis.\u201d <em>American Journal of Physical Anthropology <\/em>69 (4): 427\u2013435.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Komar, Debra A., and Jane E. Buikstra. 2008. <em>Forensic Anthropology: Contemporary Theory and Practice<\/em>. New York: Oxford University Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Langley, Natalie R., Alice F. Gooding, and MariaTeresa Tersigni-Tarrant. 2017. \u201cAge Estimation Methods.\u201d In <em>Forensic Anthropology: A Comprehensive Introduction, Second Edition<\/em>, edited by Natalie R. Langley and MariaTeresa A. Tersigni-Tarrant, 175\u2013191. Boca Raton, FL: CRC Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Lovell, Nancy C. 1997. \u201cTrauma Analysis in Paleopathology.\u201d <em>Yearbook of Physical Anthropology<\/em> 104 (S25): 139\u2013170.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Native American Graves Protection and Repatriation Act (NAGPRA) 1990 (25 U.S. Code 3001 et seq.)<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">NIST (National Institute of Standards and Technology). N.d. \u201cThe Organization of Scientific Area Committees for Forensic Science.\u201d Accessed April 18, 2023. <a class=\"rId120\" href=\"https:\/\/www.nist.gov\/topics\/organization-scientific-area-committees-forensic-science\">https:\/\/www.nist.gov\/topics\/organization-scientific-area-committees-forensic-science<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ousley, Stephen. 1995. \u201cShould We Estimate Biological or Forensic Stature?\u201d <em>Journal of Forensic Sciences<\/em> 40(5): 768\u2013773.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Phenice, T. W. 1969. \u201cA Newly Developed Visual Method of Sexing the Os Pubis.\u201d <em>American Journal of Physical Anthropology<\/em> 30 (2): 297\u2013302.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Rose, Jerome C., Thomas J. Green, and Victoria D. Green. 1996. \u201cNAGPRA Is Forever: Osteology and the Repatriation of Skeletons.\u201d <em>Annual Review of Anthropology <\/em>25: 81\u2013103.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Schaefer, Maureen, Sue Black, and Louise Scheuer. <em>Juvenile Osteology: A Laboratory and Field Manua<\/em>l. 2009. San Diego: Elsevier Academic Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Schall, Jenna L., Tracy L. Rogers, and Jordan D. Deschamps-Braly. 2020. \u201cBreaking the Binary: The Identification of Trans-women in Forensic Anthropology.\u201d <em>Forensic Science International<\/em> 309: 110220. https:\/\/doi.org\/10.1016\/j.forsciint.2020.110220.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2010a. \u201cPersonal Identification.\u201d Last modified June 3, 2010. <a class=\"rId121\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_personal_identification.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_personal_identification.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2010b. \u201cSex Assessment.\u201d Last modified June 3, 2010. <a class=\"rId122\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_sex_assessment.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_sex_assessment.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2011. \u201cTrauma Analysis.\u201d Last modified May 27, 2011. <a class=\"rId123\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_trauma.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_trauma.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2012. \u201cStature Estimation.\u201d Last modified August 2, 2012. <a class=\"rId124\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_stature_estimation.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_stature_estimation.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2013. \u201cAge Estimation.\u201d Last modified January 22, 2013. <a class=\"rId125\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_age_estimation.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_age_estimation.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Soler, Angela, and Jared S. Beatrice. 2018. \u201cExpanding the Role of Forensic Anthropology in Humanitarian Crisis: An Example from the USA-Mexico Border. In <em>Sociopolitics of Migrant Death and Repatriation: Perspectives from Forensic Science<\/em>, edited by Krista E. Latham and Alyson J. O\u2019Daniel, 115\u2013128. New York: Springer.<\/p>\n<p class=\"import-Normal\">Soler, Angela, Robin Reineke, Jared Beatrice, and Bruce E. Anderson. 2019. \u201cEtched in Bone: Embodied Suffering in the Remains of Undocumented Migrants.\u201d <em>In<\/em> <em>The Border and Its Bodies: The Embodiment of Risk along the U.S.-M\u00e9xico Line<\/em>, edited by Thomas E. Sheridan and Randall H. McGuire, 173\u2013207. Tucson: University of Arizona Press.<\/p>\n<p class=\"import-Normal\">Stull, Kyra E., Eric J. Bartelink, Alexandra R. Klales, Gregory E. Berg, Michael W. Kenyhercz, Erica N. L\u2019Abb\u00e9, Matthew C. Go, et al.. 2021. \u201cCommentary on: Bethard JD, DiGangi EA. Letter to the Editor\u2014Moving Beyond a Lost Cause: Forensic Anthropology and Ancestry Estimates in the United States. J Forensic Sci. 2020;65(5):1791\u20132. doi: 10.1111\/1556-4029.14513.\u201d <em>Journal of Forensic Sciences <\/em>66 (1): 417\u2013420.<\/p>\n<p class=\"import-Normal\">Tallman, Sean D., Caroline D. Kincer, and Eric D. Plemons. 2022. \u201cCentering Transgender Individuals in Forensic Anthropology and Expanding Binary Sex Estimation in Casework and Research.\u201d Special issue, \u201cDiversity and Inclusion,\u201d <em>Forensic Anthropology<\/em> 5 (2): 161\u2013180.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Tersigni-Tarrant, MariaTeresa A., and Natalie R. Langley. 2017. \u201cHuman Osteology.\u201d In <em>Forensic Anthropology: A Comprehensive Introduction, Second Edition<\/em>, edited by Natalie R. Langley and MariaTeresa A. Tersigni-Tarrant, 81\u2013109. Boca Raton, FL: CRC Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ubelaker, Douglas H. 2018. \u201cA History of Forensic Anthropology.\u201d Special issue, \u201cCentennial Anniversary Issue of AJPA,\u201d <em>American Journal of Physical Anthropology<\/em> 165 (4): 915\u2013923.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">White, Tim D., and Pieter A. Folkens. 2005. <em>The Human Bone Manual<\/em>. Burlington, MA: Elsevier Academic Press.<\/p>\n<p class=\"import-Normal\">Winburn, Allysha P., and Bridget Algee-Hewitt. 2021. \u201cEvaluating Population Affinity Estimates in Forensic Anthropology: Insights from the Forensic Anthropology Database for Assessing Methods Accuracy (FADAMA).\u201d <em>Journal of Forensic Sciences<\/em> 66 (4): 1210\u20131219.<\/p>\n<p class=\"import-Normal\">Winburn, Allysha Powanda, Sarah Kiley Schoff, and Michael W. Warren. 2016. \u201cAssemblages of the Dead: Interpreting the Biocultural and Taphonomic Signature of Afro- Cuban Palo Practice in Florida.\u201d <em>Journal of African Diaspora Archaeology and Heritage <\/em>5 (1): 1\u201337.<\/p>\n<\/div>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1780\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1780\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\">Ashley Kendell, Ph.D., California State University, Chico<\/p>\n<p class=\"import-Normal\">Alex Perrone, M.A., M.S.N, R.N., P.H.N., Butte Community College<\/p>\n<p class=\"import-Normal\">Colleen Milligan, Ph.D., California State University, Chico<\/p>\n<p class=\"import-Normal\"><em>This chapter is a revision from \"<\/em><a class=\"rId7\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\"><em>Chapter 15: Bioarchaeology and Forensic Anthropology<\/em><\/a><em>\u201d by Ashley Kendell, Alex Peronne, and Colleen Milligan. In <\/em><a class=\"rId8\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, first edition<\/em><\/a><em>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under <\/em><a class=\"rId9\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\"><em>CC BY-NC 4.0<\/em><\/a><em>. <\/em><\/p>\n<p class=\"import-Normal\"><strong>Content Warning and Disclaimer:<\/strong> This chapter includes images of human remains as well as discussions centered on human skeletal analyses. All images are derived from casts, sketches, nonhuman skeletal material, as well as non-Indigenous skeletal materials curated within the CSU, Chico Human Identification Lab, and the Hartnett-Fulginiti donated skeletal collection.<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\"><span style=\"color: #ffffff\">Learning Objectives<\/span><\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Define forensic anthropology as a subfield of biological anthropology.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Describe the seven steps carried out during skeletal analysis.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Outline the four major components of the biological profile.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Contrast the four categories of trauma.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Explain how to identify the different taphonomic agents that alter bone.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Discuss ethical considerations for forensic anthropology.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1214\">Forensic anthropology<\/a><\/strong> is a subfield of biological anthropology and an applied area of anthropology. Forensic anthropologists use skeletal analysis to gain information about humans in the present or recent past, then they apply this information within a medicolegal context. This means that forensic anthropologists specifically conduct their analysis on recently deceased individuals (typically within the last 50 years) as part of investigations by law enforcement. Forensic anthropologists can assist law enforcement agencies in several different ways, including aiding in the identification of human remains whether they are complete, fragmentary, burned, scattered, or decomposed. Additionally, forensic anthropologists can help determine what happened to the deceased at or around the time of death as well as what processes acted on the body after death (e.g., whether the remains were scattered by animals, whether they were buried in the ground, or whether they remained on the surface as the soft tissue decomposed).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Many times, because of their expertise in identifying human skeletal remains, forensic anthropologists are called to help with outdoor search-and-recovery efforts, such as locating remains scattered across the surface or carefully excavating and documenting buried remains. In other cases, forensic anthropologists recover remains after natural disasters or accidents, such as fire scenes, and can help identify whether each bone belongs to a human or an animal. Forensic anthropology spans a wide scope of contexts involving the law, including incidences of mass disasters, genocide, and war crimes.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">A point that can be somewhat confusing for students is that although the term <em>forensic<\/em> is included in this subfield of biological anthropology, there are many forensic techniques that are not included in the subfield. Almost exclusively, forensic anthropology deals with skeletal analysis. While this can include the comparison of antemortem (before death) and postmortem (after death) radiographs to identify whether remains belong to a specific person, or using photographic superimposition of the cranium, it does not include analyses beyond the skeleton. For example, blood-spatter analysis, DNA analysis, fingerprints, and material evidence collection do not fall under the scope of forensic anthropology.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">So, what can forensic anthropologists glean from bones alone? Forensic anthropologists can address a number of questions about a human individual based on their skeletal remains. Some of those questions are as follows: How old was the person? Was the person biologically male or female? How tall was the person? What happened to the person at or around their time of death? Were they sick? The information from the skeletal analysis can then be matched with missing persons records, medical records, or dental records, aiding law enforcement agencies with identifications and investigations.<\/p>\n<h2 class=\"import-Normal\">Skeletal Analysis<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropology relies on skeletal analysis to reveal information about the deceased. <span style=\"background-color: #00ffff\">The methodology and approaches outlined below are specific to the United States.<\/span> Forensic anthropological methods differ depending on the country conducting an investigation. In the United States, there are typically seven steps or questions to the process:<\/p>\n<ul>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Is it bone?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Is it human?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Is it modern or archeological?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">How many individuals are present or what is the minimum number of individuals (MNI)?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Who is it?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Is there evidence of trauma before or around the time of death?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">What happened to the remains after death?<\/li>\n<\/ul>\n<h3 class=\"import-Normal\"><strong>Is It Bone?<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">One of the most important steps in any skeletal analysis starts with determining whether or not material suspected to be bone is in fact bone. Though it goes without saying that a forensic anthropologist would only carry out analysis on bone, this step is not always straightforward. Whole bones are relatively easy to identify, but determining whether or not something is bone becomes more challenging once it becomes fragmentary. As an example, in high heat such as that seen on fire scenes, bone can break into pieces. During a house fire with fatalities, firefighters watered down the burning home. After the fire was extinguished, the sheetrock (used to construct the walls of the home) was drenched and crumbled. The crumbled sheetrock was similar in color and form to burned, fragmented bone, therefore mistakable for human remains (Figure 15.1). Forensic anthropologists on scene were able to separate the bones from the construction material, helping to confirm the presence of bone and hence the presence of individual victims of the fire. In this case, forensic anthropologists were able to recognize the anatomical and layered structure of bone and were able to distinguish it from the uniform and unlayered structure of sheetrock.<\/p>\n<p class=\"import-Normal\"><strong><img class=\"aligncenter\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/06\/image3.png\" alt=\"Long rectangular sheetrock with exposed porous surface.\" width=\"182\" height=\"208\" \/><\/strong><\/p>\n<figure style=\"width: 372px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image6-1.png\" alt=\"Two examples of sheetrock with dried or burnt surfaces.\" width=\"372\" height=\"210\" \/><figcaption class=\"wp-caption-text\">Figure 15.1: Burned sheetrock used as building material appears similar to human bone but can be differentiated by the fact that it is the same density throughout. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Example of burned sheetrock (Figure 15.1)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">As demonstrated by the example above, both the macrostructure (visible with the naked eye) and microstructure (visible with a microscope) of bone are helpful in bone identification. Bones are organs in the body made up of connective tissue. The connective tissue is hardened by a mineral deposition, which is why bone is rigid in comparison to other connective tissues such as cartilage (Tersigni-Tarrant and Langley 2017, 82\u201383; White and Folkens 2005, 31). In a living body, the mineralized tissue does not make up the only component of bone\u2014there are also blood, bone marrow, cartilage, and other types of tissues. However, in dry bone, two distinct layers of the bone are the most helpful for identification. The outer layer is made up of densely arranged osseous (bone) tissue called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1216\">compact (cortical) bone<\/a><\/strong>. The inner layer is composed of much more loosely organized, porous bone tissue whose appearance resembles that of a sponge, hence the name <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1218\"><strong>spongy (trabecular) bone<\/strong><\/a>. Knowing that most bone contains both layers helps with the macroscopic identification of bone (Figures 15.2, 15.3). For example, a piece of coconut shell might look a lot like a fragment of a human skull bone. However, closer inspection will demonstrate that coconut shell only has one very dense layer, while bone has both the compact and spongy layers.<\/p>\n<figure style=\"width: 380px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image27-1.png\" alt=\"Drawing showing thick exterior compact bone and porous internal cortical bone.\" width=\"380\" height=\"371\" \/><figcaption class=\"wp-caption-text\">Figure 15.2: Cross section of human long bone with compact and cortical bone layers visible. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Cross section of human long bone (Figure 15.2)<\/a> original to<a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"> Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p>&nbsp;<\/p>\n<figure style=\"width: 364px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image25-2.png\" alt=\"Cranial bone cross section called a periosteum with spongy bone (diploe) and compact bone labeled. Compact bone is a thin slice at the top and bottom and is smooth and hard. Spongy bone is in the middle and has irregular holes and indentations throughout. \" width=\"364\" height=\"184\" \/><figcaption class=\"wp-caption-text\">Figure 15.3: Cranial anatomy is slightly different as compared to that of a long bone in cross section. The compact (cortical) bone layers sandwich the spongy (trabecular) bone. One layer of compact bone forms the very outer surface of the skull and the other lines the internal surface of the skull. Credit: <a href=\"https:\/\/cnx.org\/contents\/FPtK1zmh@6.27:kwbeYj9S@3\/Bone-Structure\">Anatomy of a Flat Bone (Anatomy &amp; Physiology, Figure 6.3.3)<\/a> by<a href=\"https:\/\/openstax.org\/\"> OpenStax<\/a> is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/\"> CC BY 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The microscopic identification of bone relies on knowledge of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1220\">osteons<\/a><\/strong>, or bone cells (Figure 15.4). Under magnification, bone cells are visible in the outer, compact layer of bone. The bone cells are arranged in a concentric pattern around blood vessels for blood supply. The specific shape of the cells can help differentiate, for example, a small piece of PVC (white plastic) pipe from a human bone fragment (Figure 15.5).<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 340px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image14-3.png\" alt=\"Microscope image showing clustered osteons. Each has many rings and a dark center.\" width=\"340\" height=\"218\" \/><figcaption class=\"wp-caption-text\">Figure 15.4: Bone microstructure (osteons). Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Bone_(248_12)_Bone_cross_section.jpg\">Bone (248 12) Bone cross section<\/a> by <a href=\"https:\/\/cs.wikipedia.org\/wiki\/Josef_Reischig\">Doc. RNDr. Josef Reischig, CSc.<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p>&nbsp;<\/p>\n<figure style=\"width: 332px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image7-1.png\" alt=\"Flat, white section of PVC. Edges are broken and surface rough.\" width=\"332\" height=\"268\" \/><figcaption class=\"wp-caption-text\">Figure 15.5: Fragments of plastic PVC pipe, such as those seen in this photo, may be mistaken for human bone. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Example of PVC pipe<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Is It Human?<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Once it has been determined that an object is bone, the next logical step is to identify whether the bone belongs to a human or an animal. Forensic anthropologists are faced with this question in everyday practice because human versus nonhuman bone identification is one of the most frequent requests they receive from law enforcement agencies.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">There are many different ways to distinguish human versus nonhuman bone. The morphology (the shape\/form) of human bone is a good place for students to start. Identifying the 206 bones in the adult human skeleton and each bone\u2019s distinguishing features (muscle attachment sites, openings and grooves for nerves and blood vessels, etc.) is fundamental to skeletal analysis.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Nevertheless, there are many animal bones and human bones that look similar. For example, the declawed skeleton of a bear paw looks a lot like a human hand, pig molars appear similar to human molars, and some smaller animal bones might be mistaken for those of an infant. To add to the confusion, fragmentary bone may be even more difficult to identify as human or nonhuman. However, several major differences between human and nonhuman vertebrate bone help distinguish the two.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropologists pay special attention to the density of the outer, compact layer of bone in both the cranium and in the long bones. Human cranial bone has three distinctive layers. The spongy bone is sandwiched between the outer (ectocranial) and inner (endocranial) compact layers. In most other mammals, the distinction between the spongy and compact layers is not always so definite. Secondly, the compact layer in nonhuman mammal long bones can be much thicker than observed in human bone. Due to the increased density of the compact layer, nonhuman bone tends to be heavier than human bone (Figure 15.6).<\/p>\n<figure style=\"width: 399px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image11-2.png\" alt=\"Ring-like cross section of bone.\" width=\"399\" height=\"266\" \/><figcaption class=\"wp-caption-text\">Figure 15.6: The compact layer of this animal bone is very thick, with almost no spongy bone visible. Compare with Figure 15.2 to visualize the difference in structure between human and nonhuman bone. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Animal bone cross section (Figure 15.6)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The size of a bone can also help determine whether it belongs to a human. Adult human bones are larger than subadult or infant bones. However, another major difference between human adult bones and those of a young individual or infant human can be attributed to development and growth of the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1222\">epiphyses<\/a><\/strong> (ends of the bone). The epiphyses of human subadult bones are not fused to the shaft (Figure 15.7). Therefore, if a bone is small and it is suspected to belong to a human subadult or infant, the epiphyses would not be fused. Many small animal bones appear very similar in form compared to adult human bones, but they are much too small to belong to an adult human. Yet they can be eliminated as subadult or infant bones if the epiphyses are fused to the shaft.<\/p>\n<figure style=\"width: 288px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image13-3.png\" alt=\"X-ray image of child\u2019s ankle.\" width=\"288\" height=\"412\" \/><figcaption class=\"wp-caption-text\">Figure 15.7: An x-ray of a subadult\u2019s ankle with the epiphyses of the tibia and fibula visible. The gap between the shaft of the bone and the end of the bone (epiphysis) is the location of the growth plate. Therefore, the growth plate gap is what separates the shafts from the epiphyses in the image. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Tib_fib_growth_plates.jpg\">Tib fib growth plates<\/a> by <a href=\"https:\/\/en.wikipedia.org\/wiki\/User:Gilo1969\">Gilo1969<\/a> at <a href=\"https:\/\/en.wikipedia.org\/wiki\/\">English Wikipedia<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/3.0\/legalcode\">CC BY 3.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Is It Modern or Archaeological? <\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropologists work with modern cases that fall within the scope of law enforcement investigations. Accordingly, it is important to determine whether discovered human remains are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1224\">archaeological<\/a> <\/strong>or forensic in nature. Human remains that are historic are considered archeaological. The scientific study of human remains from archaeological sites is called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1226\">bioarchaeology<\/a><\/strong>.<\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Dig Deeper: Bioarchaeology<\/h2>\n<p class=\"import-Normal\">For readers who are interested in the sister subfield of bioarchaeology, which studies human remains and material culture from the past, please refer to chapter 8 of <em>Bioarchaeology: Interpreting Human Behavior from Skeletal Remains,<\/em> in <em>TRACES: An Open Invitation to Archaeology<\/em> (Blatt, Michael, and Bright forthcoming).<\/p>\n<\/div>\n<p>A forensic anthropologist should begin their analysis by reviewing the context in which the remains were discovered. This will help them understand a great deal about the remains, including determining whether they are archaeological or forensic in nature as well as considering legal and ethical issues associated with the collection, analysis, and storage of human remains (see \u201cEthics and Human Rights\u201d section of this chapter for more information).<\/p>\n<figure style=\"width: 403px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image10-3.png\" alt=\"Four teeth in a person\u2019s mouth. First molar with silver filling.\" width=\"403\" height=\"303\" \/><figcaption class=\"wp-caption-text\">Figure 15.8: A human tooth with a filling. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Filling.jpg#filehistory\">Filling<\/a> by Kauzio has been designated to the <a href=\"https:\/\/creativecommons.org\/share-your-work\/public-domain\/cc0\/\">public domain (CC0)<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The \u201ccontext\u201d refers to the relationship the remains have to the immediate area in which they were found. This includes the specific place where the remains were found, the soil or other organic matter immediately surrounding the remains, and any other objects or artifacts in close proximity to the body. For example, imagine that a set of remains has been located during a house renovation. The remains are discovered below the foundation. Do the remains belong to a murder victim? Or was the house built on top of an ancient burial ground? Observing information from the surroundings can help determine whether the remains are archaeological or modern. How long ago was the foundation of the house erected? Are there artifacts in close proximity to the body, such as clothing or stone tools? These are questions about the surroundings that will help determine the relative age of the remains.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Clues directly from the skeleton may also indicate whether the remains are archaeological or modern. For example, tooth fillings can suggest that the individual was alive recently (Figure 15.8). In fact, filling material has changed over the decades, so the specific type of material used to fix a cavity can be matched with specific time periods. Gold was used in dental work in the past, but more recently composite (a mixture of plastic and fine glass) fillings have become more common.<\/p>\n<h3><strong>How <\/strong><strong>Many Individuals Are Present?<\/strong><\/h3>\n<h4 class=\"import-Normal\"><em>What Is MNI?<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Another assessment that an anthropologist can perform is the calculation of the number of individuals in a mixed burial assemblage. Because not all burials consist of a single individual, it is important to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1268\">burial assemblage<\/a><\/strong> be able to estimate the number of individuals in a forensic context. Quantification of the number of individuals in a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1524\">burial assemblage<\/a><\/strong> can be done through the application of a number of methods, including the following: the Minimum Number of Individuals (MNI), the Most Likely Number of Individuals (MLNI), and the Lincoln Index (LI). The most commonly used method in biological anthropology, and the focus of this section, is determination of the MNI.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The MNI presents \u201cthe minimum estimate for the number of individuals that contributed to the sample\u201d (Adams and Konigsberg 2008, 243). Many methods of calculating MNI were originally developed within the field of zooarchaeology for use on calculating the number of individuals in faunal or animal assemblages (Adams and Konigsberg 2008, 241). What MNI calculations provide is a lowest possible count for the total number of individuals contributing to a skeletal assemblage. Traditional methods of calculating MNI include separating a skeletal assemblage into categories according to the individual bone and the side the bone comes from and then taking the highest count per category and assigning that as the minimum number (Figure 15.9).<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 664px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image28-3.png\" alt=\"Many bone portions laying on individual plastic bags on a table.\" width=\"664\" height=\"441\" \/><figcaption class=\"wp-caption-text\">Figure 15.9: Skeletal elements from a commingled faunal assemblage. Credit: Commingled animal remains from Eden-Farson Pre-Contact site in southwest Wyoming by Matt O\u2019Brien original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Why Calculate MNI?<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In a forensic context, the determination of MNI is most applicable in cases of mass graves, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1232\">commingled burials<\/a><\/strong>, and mass fatality incidents. The term <em>commingled<\/em> is applied to any burial assemblage in which individual skeletons are not separated into separate burials. As an example, the authors of this chapter have observed commingling of remains resulting from mass fatality wildfire events. Commingled remains may also be encountered in events such as a plane or vehicle crash. It is important to remember that in any forensic context, MNI should be referenced and an MNI of one should be substantiated by the fact that there was no repetition of elements associated with the case.<\/p>\n<h3 class=\"import-Normal\"><strong>Constructing the Biological Profile<\/strong><\/h3>\n<h4 class=\"import-Normal\"><em>Who Is It?<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">\u201cWho is it?\u201d is one of the first questions that law enforcement officers ask when they are faced with a set of skeletal remains. To answer this question, forensic anthropologists construct a biological profile (White and Folkens 2005, 405). A <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1228\">biological profile<\/a> <\/strong>is an individual\u2019s identifying characteristics, or biological information, which include the following: biological sex, age at death, stature, population affinity, skeletal variation, and evidence of trauma and pathology.<\/p>\n<h4 class=\"import-Normal\"><em>Assessing Biological Sex <\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Assessment of biological sex is often one of the first things considered when establishing a biological profile because several other parts, such as age and stature estimations, rely on an assessment of biological sex to make the calculations more accurate.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Assessment of biological sex focuses on differences in both morphological (form or structure) and metric (measured) traits in individuals. When assessing morphological traits, the skull and the pelvis are the most commonly referenced areas of the skeleton. These differences are related to sexual dimorphism usually varying in the amount of robusticity seen between males and females. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1230\">Robusticity<\/a> <\/strong>deals with strength and size; it is frequently used as a term to describe a large size or thickness. In general, males will show a greater degree of robusticity than females. For example, the length and width of the mastoid process, a bony projection located behind the opening for the ear, is typically larger in males. The mastoid process is an attachment point for muscles of the neck, and this bony projection tends to be wider and longer in males. In general, cranial features tend to be more robust in males (Figure 15.10).<\/p>\n<figure style=\"width: 601px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image26-3.png\" alt=\"Front and side images of a male (left) and female (right) cranium.\" width=\"601\" height=\"632\" \/><figcaption class=\"wp-caption-text\">Figure 15.10: Anterior and lateral view of a male and female cranium. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Anterior and lateral view of a male and female cranium (Figure 15.10)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropo logy<\/a> by Ashley Kendell is a collective work under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Includes <a href=\"https:\/\/boneclones.com\/product\/modern-human-asian-female-skull-BC-149\/category\/all-human-skulls\/human-anatomy\">Human Female Asian Skull<\/a> and <a href=\"https:\/\/boneclones.com\/product\/human-asian-male-skull-BC-016\/category\/all-human-skulls\/human-anatomy\">Human Male Asian Skull<\/a> by <a href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a>, used by permission.]<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">When considering the pelvis, the features associated with the ability to give birth help distinguish females from males. During puberty, estrogen causes a widening of the female pelvis to allow for the passage of a baby. Several studies have identified specific features or bony landmarks associated with the widening of the hips, and this section will discuss one such method. The Phenice Method (Phenice 1969) is traditionally the most common reference used to assess morphological characteristics associated with sex. The Phenice Method specifically looks at the presence or absence of (1) a ventral arc, (2) the presence or absence of a subpubic concavity, and (3) the width of the medial aspect of the ischiopubic ramus (Figure 15.11). When present, the ventral arc, a ridge of bone located on the ventral surface of the pubic bone, is indicative of female remains. Likewise the presence of a subpubic concavity and a narrow medial aspect of the ischiopubic ramus is associated with a female sex estimation. Assessments of these features, as well as those of the skull (when both the pelvis and skull are present), are combined for an overall estimation of sex.<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 1603px\" class=\"wp-caption alignnone\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image29-3.png\" alt=\"Male and female os coxae (anterior portions).\" width=\"1603\" height=\"582\" \/><figcaption class=\"wp-caption-text\">Figure 15.11: Features associated with the Phenice Method. Images derived from CSU-HIL donated skeletal collection. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Features associated with the Phenice Method (Figure 15.11)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Colleen Milligan is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Metric analyses are also used in the estimation of sex. Measurements taken from every region of the body can contribute to estimating sex through statistical approaches that assign a predictive value of sex. These approaches can include multiple measurements from several skeletal elements in what is called multivariate (multiple variables) statistics. Other approaches consider a single measurement, such as the diameter of the head of the femur, of a specific element in a univariate (single variable) analysis (Berg 2017, 152\u2013156).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">It is important to note that, although forensic anthropologists usually begin assessment of biological profile with biological sex, there is one major instance in which this is not appropriate. The case of two individuals found in California, on July 8, 1979, is one example that demonstrates the effect age has on the estimation of sex. The identities of the two individuals were unknown; therefore, law enforcement sent them to a lab for identification. A skeletal analysis determined that the remains represented one adolescent male and one adolescent female, both younger than 18 years of age. This information did not match with any known missing children at the time.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In 2015, the cold case was reanalyzed, and DNA samples were extracted. The results indicated that the remains were actually those of two girls who went missing in 1978. The girls were 15 years old and 14 years old at the time of death. It is clear that the 1979 results were incorrect, but this mistake also provides the opportunity to discuss the limitations of assessing sex from a subadult skeleton.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Assessing sex from the human skeleton is based on biological and genetic traits associated with females and males. These traits are linked to differences in sexual dimorphism and reproductive characteristics between females and males. The link to reproductive characteristics means that most indicators of biological sex do not fully manifest in prepubescent individuals, making estimations of sex unreliable in younger individuals (SWGANTH 2010b). This was the case in the example of the 14-year-old girl. When examined in 1979, her remains were misidentified as male because she had not yet fully developed female pelvic traits.<\/p>\n<h4 class=\"import-Normal\"><em>Sex vs. Gender<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Biological sex is a different concept than <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1234\">gender<\/a><\/strong>. While biological anthropologists can estimate sex from the skeleton, estimating an individual\u2019s gender would require a greater context because gender is defined culturally rather than biologically. Take, for example, an individual who identifies as transgender. This individual has a gender identity that is different from their biological sex. The gender identity of any individual depends on factors related to self-identification, situation or context, and cultural factors. <span style=\"background-color: #00ffff\">While in the U.S<\/span>. we have historically thought of sex and gender as binary concepts (male or female), many cultures throughout the world recognize several possible gender identities. In this sense, gender is seen as a continuous or fluid variable rather than a fixed one.<\/p>\n<p class=\"import-Normal\">Historically, forensic anthropologists have used a binary construct to categorize human skeletal remains as either male or female (with the accompanying categories of probable male, probable female, and indeterminate). In the case of transgender and gender nonconforming individuals, the binary approach to sex assessment may delay or hinder identification efforts (Buchanan 2014; Schall, Rogers, and Deschamps-Braly 2020; Tallman, Kincer, and Plemons 2021). As such, many forensic anthropologists have begun to address the inherent problems associated with a binary approach to sex identification and to explore ways of assessing social identity and self-identified gender using skeletal remains and forensic context.<\/p>\n<p class=\"import-Normal\">For the duration of this section, the term <em>transgender<\/em> refers to individuals whose gender identity differs from the sex assigned at birth (Schall, Rogers, and Deschamps-Braly 2020:2). Transgender individuals transition from one gender binary to another, such as male-to-female (MTF) or female-to-male (FTM). While many of the gender-affirming procedures available to trans and gender-nonconforming individuals are focused on soft tissue modifications (e.g., breast augmentation, genital reconstruction, hormone therapies, etc.), there are a number of gender-affirmation surgeries that do leave a permanent record on the skeleton. Generally speaking, FTM transgender people are reported to undergo fewer surgical procedures than do MTF transgender people (Buchanan 2014). The discussion below focuses on Facial Feminization Surgery (FFS), which leaves a permanent record on the human skeleton that may be used to help make an identification.<\/p>\n<p class=\"import-Normal\">FFS refers to a combination of procedures focused on sexually dimorphic features of the face, with the intent of transforming typically male facial features into more feminine forms. Facial Feminization Surgery procedures were developed by Dr. Douglas Ousterhout, a San Francisco based cranio-maxillofacial surgeon, in the mid-1980s (Schall, Rogers, and Deschamps-Braly 2020:2). FFS can include one or a combination of the following: hairline lowering, forehead reduction and contouring, brow lift, reduction rhinoplasty, cheek enhancement, lift lift, lip filling, chin contouring, jaw contouring, and\/or tracheal shave (Buchanan 2014; Schall, Rogers, and Deschamps-Braly 2020:2). Of the procedures outlined previously, four are known to directly affect the facial skeleton: forehead contouring, rhinoplasty, chin contouring, and jaw contouring (Buchanan 2014; Schall, Rogers, and Deschamps-Braly 2020:2).<\/p>\n<p class=\"import-Normal\">Because FFS procedures have been widely documented in the medical (and more recently the forensic anthropological) literature, there are a number of indicators that a forensic anthropologist can use to make more informed evaluations of gender, including evidence of bone remodeling in sexually dimorphic regions of the skull (e.g., forehead, chin, jawline), as well as the presence of plates, pins, or other surgical hardware that may be evidence of FFS (Buchanan 2014; Schall, Rogers, and Deschamps-Braly 2020; Tallman, Kincer, and Plemons 2021). Additionally, some forensic anthropologists suggest cautiously integrating contextual information from the scene, such as personal effects, material evidence, and recovery scene information, into their evaluation of an individual\u2019s social identity (Beatrice and Soler 2016; Birkby, Fenton, and Anderson 2008; Soler and Beatrice 2018; Soler et al. 2019; Tallman, Kincer, and Plemons 2021; Winburn, Schoff, and Warren 2016). The ultimate goal of many skeletal analyses is to make a positive identification on a set of unidentified remains.<\/p>\n<h4 class=\"import-Normal\"><em>Assessment <\/em><em>of Population Affinity<\/em><\/h4>\n<p>In an effort to combat the erroneous assumptions tied to the race concept, forensic anthropologists have attempted to reframe this component of the biological profile. The term <em>race<\/em> is no longer used in casework and teaching. Historically, the word <em>ancestry<\/em> is and was deemed a more appropriate way to describe an individual\u2019s phenotype. However, in more recent years, forensic anthropologists have begun using the term <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1236\">population affinity<\/a><\/strong><em>, <\/em>recognizing that we are basing our analysis on the similarities we see based on the reference samples we have available (Winburn and Algee-Hewitt 2021). An important note here is that it is possible to hinder identifications and harm individuals when tools like estimations of population affinity are misapplied, misinterpreted, or misused. For this reason, the field of forensic anthropology has ongoing conversations about the appropriateness of this analysis in the biological profile (Bethard and DiGangi 2020; Stull et al. 2021).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">We use the term <em>population affinity<\/em> to refer to the variation seen among modern populations\u2014variation that is both genetic and environmentally driven. The word <em>affinity<\/em> refers to similarities or relationships between individuals. As forensic anthropologists, we compare an unknown individual to multiple reference groups and look for the degree of similarity in observable traits with those groups. As noted previously, population affinity can aid law enforcement in their identification of missing persons or unknown skeletal remains.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Within the field of anthropology, the estimation of population affinity has a contentious history, and early attempts at classification were largely based on the erroneous assumption that an individual\u2019s <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1238\">phenotype <\/a><\/strong> (outward appearance) was correlated with their innate intelligence and abilities (see Chapter 13 for a more in-depth discussion of the history of the race concept). The use of the term <em>race<\/em> is deeply embedded in the social context of the United States. In any other organism\/living thing, groups divided according to the biological race concept would be defined as a separate subspecies. The major issue with applying the biological race concept to humans is that there are not enough differences between any two populations to separate on a genetic basis. In other words, <em>biological races do not exist in human populations. <\/em>However, the concept of race has been perpetuated and upheld by sociocultural constructs of race.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The conundrum for forensic anthropologists is the fact that while races do not exist on a biological level, we still socially recognize and categorize individuals based on their phenotype. Clearly, our phenotype is an important factor in not only how we are viewed by others but also how we identify ourselves. It is also a commonly reported variable. Often labeled as \u201crace,\u201d we are asked to report how we self-identify on school applications, government identification, surveys, census reports, and so forth. It follows then that when a person is reported missing, the information commonly collected by law enforcement and sometimes entered into a missing person\u2019s database includes their age, biological sex, stature, and \u201crace.\u201d Therefore, the more information a forensic anthropologist can provide regarding the individual\u2019s physical characteristics, the more he or she can help to narrow the search.<\/p>\n<p class=\"import-Normal\">As an exercise, create a list of all of the women you know who are between the ages of 18 and 24 and approximately 5\u2019 4\u201d to 5\u2019 9\u201d tall. You probably have several dozen people on the list. Now, consider how many females you know who are between the ages of 18 and 24, are approximately 5\u2019 4\u201d to 5\u2019 9\u201d tall, and are Vietnamese. Your list is going to be significantly shorter. That\u2019s how missing persons searches go as well. The more information you can provide regarding a decedent\u2019s phenotype, the fewer possible matches law enforcement are left to investigate. This is why population affinity has historically been included as a part of the biological profile.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Traditionally, population affinity was accomplished through a visual inspection of morphological variants of the skull (morphoscopics). These methods focused on elements of the facial skeleton, including the nose, eyes, and cheek bones. However, in an effort to reduce subjectivity, nonmetric cranial traits are now assessed within a statistical framework to help anthropologists better interpret their distribution among living populations (Hefner and Linde 2018). Based on the observable traits, a macromorphoscopic analysis will allow the practitioner to create a statistical prediction of geographic origin. In essence, forensic anthropologists are using human variation in the estimation of geographic origin, by referencing documented frequencies of nonmetric skeletal indicators or macromorphoscopic traits.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Population affinity is also assessed through metric analyses. The computer program Fordisc is an anthropological tool used to estimate different components of the biological profile, including ancestry, sex, and stature. When using Fordisc, skeletal measurements are input into the computer software, and the program employs multivariate statistical classification methods, including discriminant function analysis, to generate a statistical prediction for the geographic origin of unknown remains based on the comparison of the unknown to the reference samples in the software program. Fordisc also calculates the likelihood of the prediction being correct, as well as how typical the metric data is for the assigned group.<\/p>\n<h4 class=\"import-Normal\"><em>Estimating Age-at-Death<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Estimating age-at-death from the skeleton relies on the measurement of two basic physiological processes: (1) growth and development and (2) degeneration (or aging). From fetal development on, our bones and teeth grow and change at a predictable rate. This provides for relatively accurate age estimates. After our bones and teeth cease to grow and develop, they begin to undergo structural changes, or degeneration, associated with aging. This does not happen at such predictable rates and, therefore, results in less accurate or larger age-range estimations.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">During growth and development stages, two primary methods used for estimations of age of subadults (those under the age of 18) are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1240\">epiphyseal union<\/a><\/strong> and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1244\">dental development.<\/a><\/strong> Epiphyseal union<strong> (<\/strong>or <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1242\">epiphyseal fusion<\/a><\/strong>) refers to the appearance and closure of the epiphyseal plates between the primary centers of growth in a bone and the subsequent centers of growth (see Figure 15.7). Prior to complete union, the cartilaginous area between the primary and secondary centers of growth is also referred to as the growth plates (Schaefer, Black, and Scheuer 2009). Different areas of the skeleton have documented differences in the appearance and closure of epiphyses, making this a reliable method for aging subadult remains (SWGANTH 2013).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">As an example of its utility in the identification process, epiphyseal development was used to identify two subadult victims of a fatal fire in Flint, Michigan, in February 2010. The remains represented two young girls, ages three and four. Due to the intensity of the fire, the subadult victims were differentiated from each other through the appearance of the patella, the kneecap. The patella is a bone that develops within the tendon of the quadriceps muscle at the knee joint. The patella begins to form around three to four years of age (Cunningham, Scheuer, and Black 2016, 407\u2013409). In the example above, radiographs of the knees showed the presence of a patella in the four-year-old girl and the absence of a clearly discernible patella in the three-year-old.<\/p>\n<figure style=\"width: 358px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image18-2.png\" alt=\"Cranial cast of child with exposed maxilla and mandible to see developing dentition.\" width=\"358\" height=\"358\" \/><figcaption class=\"wp-caption-text\">Figure 15.12: Dental development in a subadult. Credit: <a href=\"https:\/\/boneclones.com\/product\/5-year-old-human-child-skull-with-mixed-dentition-exposed-BC-189\">5-year-old Human Child Skull with Mixed Dentition Exposed<\/a> by <a href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a> is used by permission and available here is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Dental development begins during fetal stages of growth and continues until the complete formation and eruption of the adult third molars (if present). The first set of teeth to appear are called deciduous or baby teeth. Individuals develop a total of 20 deciduous teeth, including incisors, canines, and molars. These are generally replaced by adult dentition as an individual grows (Figure 15.12). A total of 32 teeth are represented in the adult dental arcade, including incisors, canines, premolars, and molars. When dental development is used for age estimations, researchers use both tooth-formation patterns and eruption schedules as determining evidence. For example, the crown of the tooth forms first followed by the formation of the tooth root. During development, an individual can exhibit a partially formed crown or a complete crown with a partially formed root. The teeth generally begin the eruption process once the crown of the tooth is complete. The developmental stages of dentition are one of the most reliable and consistent aging methods for subadults (Langley, Gooding, and Tersigni-Tarrant 2017, 176\u2013177).<\/p>\n<figure style=\"width: 403px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image22-3.png\" alt=\"Surfaces of three pubic symphyses: billowy (A) to more flat (B) to rough (C).\" width=\"403\" height=\"224\" \/><figcaption class=\"wp-caption-text\">Figure 15.13: Examples of degenerative changes to the pubic symphysis: (A) young adult; (B) middle adult; (C) old adult. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Example of the progression of degenerative changes to the pubic symphysis (Figure 15.14)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropo logy<\/a> by Ashley Kendell is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Original photos by Dr. Julie Fleischman used by permission. Pubic symphyses are curated in the Hartnett-Fulginiti donated skeletal collection. Donation and research consent was provided by next of kin.]<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Degenerative changes in the skeleton typically begin after 18 years of age, with more prominent changes developing after an individual reaches middle adulthood (commonly defined as after 35 years of age in osteology). These changes are most easily seen around joint surfaces of the pelvis, the cranial vault, and the ribs. In this chapter, we focus on the pubic symphysis surfaces of the pelvis and the sternal ends of the ribs, which show metamorphic changes from young adulthood to older adulthood. The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1262\">pubic symphysis<\/a> <\/strong>is a joint that unites the left and right halves of the pelvis. The surface of the pubic symphysis changes during adulthood, beginning as a surface with pronounced ridges (called billowing) and flattening with a more distinct rim to the pubic symphysis as an individual ages. As with all metamorphic age changes, older adults tend to develop lipping around the joint surfaces as well as a breakdown of the joint surfaces. The most commonly used method for aging adult skeletons from the pubic symphysis is the Suchey-Brooks method (Brooks and Suchey 1990; Katz and Suchey 1986). This method divides the changes seen with the pubic symphysis into six phases based on macroscopic age-related changes to the surface. Figure 15.13 provides a visual of the degenerative changes that typically occur on the pubic symphysis.<\/p>\n<figure style=\"width: 403px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image12-3.png\" alt=\"Three sternal rib ends demonstrating progressive changes that occur with age.\" width=\"403\" height=\"220\" \/><figcaption class=\"wp-caption-text\">Figure 15.14: Examples of degenerative changes to the sternal rib end: (A) young adult; (B) middle adult; (C) old adult. Images derived from CSU, Chico HIL donated skeletal collection. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Examples of degenerative changes to the sternal rib end (Figure 15.15)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The sternal end of the ribs, the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1246\">anterior <\/a><\/strong> end of the rib that connects via cartilage to the sternum, is also used in age estimations of adults. This method, first developed by M. Y. \u0130\u015fcan and colleagues, considers both the change in shape of the sternal end as well as the quality of the bone (\u0130\u015fcan, Loth, and Wright 1984; \u0130\u015fcan, Loth, and Wright 1985). The sternal end first develops a billowing appearance in young adulthood. The bone typically develops a wider and deeper cupped end as an individual ages. Older adults tend to exhibit bony extensions of the sternal end rim as attaching cartilage ossifies. Figure 15.14 provides a visual of the degenerative changes that typically occur in sternal rib ends.<\/p>\n<h4 class=\"import-Normal\"><em>Estimating Stature<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Stature, or height, is one of the most prominently recorded components of the biological profile. Our height is recorded from infancy through adulthood. Doctor\u2019s appointments, driver's license applications, and sports rosters all typically involve a measure of stature for an individual. As such, it is also a component of the biological profile nearly every individual will have on record. Bioarchaeologists and forensic anthropologists use stature estimation methods to provide a range within which an individual\u2019s biological height would fall. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1248\">Biological height<\/a> <\/strong>is a person\u2019s true anatomical height. However, the range created through these estimations is often compared to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1264\">reported stature<\/a><\/strong>, which is typically self-reported and based on an approximation of an individual\u2019s true height (Ousley 1995).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In June 2015, two men were shot and killed in Granite Bay, California, in a double homicide. Investigators were able to locate surveillance camera footage from a gas station where the two victims were spotted in a car with another individual believed to be the perpetrator in the case. The suspect, sitting behind the victims in the car, hung his right arm out of the window as the car drove away. The search for the perpetrator was eventually narrowed down to two suspects. One suspect was 5\u2019 8\u201d while the other suspect was 6\u2019 4\u201d, representing almost a foot difference in height reported stature between the two. Forensic anthropologists were given the dimensions of the car (for proportionality of the arm) and were asked to calculate the stature of the suspect in the car from measurements of the suspect\u2019s forearm hanging from the window. Approximate lengths of the bones of the forearm were established from the video footage and used to create a predicted stature range. Stature estimations from skeletal remains typically look at the correlation between the measurements of any individual bone and the overall measurement of body height. In the case above, the length of the right forearm pointed to the taller of the two suspects who was subsequently arrested for the homicide.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Certain bones, such as the long bones of the leg, contribute more to our overall height than others and can be used with mathematical equations known as regression equations. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1252\">Regression methods <\/a> <\/strong>examine the relationship between variables such as height and bone length and use the correlation between the variables to create a prediction interval (or range) for estimated stature. This method for calculating stature is the most commonly used method (SWGANTH 2012). Figure 15.15 shows the measurement of the bicondylar length of the femur for stature estimations.<\/p>\n<figure style=\"width: 584px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image2-3.png\" alt=\"A femur is measured using a wooden osteometric board.\" width=\"584\" height=\"389\" \/><figcaption class=\"wp-caption-text\">Figure 15.15: Image of measurement of the bicondylar length of the femur, often used in the estimation of living stature. Image derived from CSU, Chico HIL donated skeletal collection. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Measurement of the bicondylar length of the femur<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Identification Using Individualizing Characteristics<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">One of the most frequently requested analyses within the forensic anthropology laboratory is assistance with the identification of unidentified remains. While all components of a biological profile, as discussed above, can assist law enforcement officers and medical examiners to narrow down the list of potential identifications, a biological profile will not lead to a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1254\">positive identification<\/a><\/strong>. The term <em>positive identification<\/em> refers to a scientifically validated method of identifying previously unidentified remains. Presumptive identifications, however, are not scientifically validated; rather, they are based on circumstances or scene context. For example, if a decedent is found in a locked home with no evidence of forced entry but the body is no longer visually identifiable, it may be presumed that the remains belong to the homeowner. Hence, a presumptive identification.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The medicolegal system ultimately requires that a positive identification be made in such circumstances, and a presumptive identification is often a good way to narrow down the pool of possibilities. Biological profile information also assists with making a presumptive identification based on an individual\u2019s phenotype in life (e.g., what they looked like). As an example, a forensic anthropologist may establish the following components of a biological profile: white male, between the ages of 35 and 50, approximately 5\u2019 7\u201d to 5\u2019 11.\u201d While this seems like a rather specific description of an individual, you can imagine that this description fits dozens, if not hundreds, of people in an urban area. Therefore, law enforcement can use the biological profile information to narrow their pool of possible identifications to include only white males who fit the age and height outlined above. Once a possible match is found, the decedent can be identified using a method of positive identification.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Positive identifications are based on what we refer to as individualizing traits or characteristics, which are traits that are unique at the individual level. For example, brown hair is not an individualizing trait as brown is the most common hair color in the U.S. But, a specific pattern of dental restorations or surgical implants can be individualizing, because it is unlikely that you will have an exact match on either of these traits when comparing two individuals.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">A number of positive methods are available to forensic anthropologists, and for the remainder of this section we will discuss the following methods: comparative medical and dental radiography and identification of surgical implants.<\/p>\n<figure style=\"width: 165px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image17-3.png\" alt=\"Radiograph of skull with frontal sinuses visible.\" width=\"165\" height=\"182\" \/><figcaption class=\"wp-caption-text\">Figure 15.16: Example of the unique shape of the frontal sinus. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Frontal_bone_sinuses.jpg\">Frontal bone sinuses<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Alex_Khimich\">Alex Khimich<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Comparative medical and dental radiography is used to find consistency of traits when comparing antemortem records (medical and dental records taken during life) with images taken postmortem (after death). Comparative medical radiography focuses primarily on features associated with the skeletal system, including trabecular pattern (internal structure of bone that is honeycomb in appearance), bone shape or cortical density (compact outer layer of bone), and evidence of past trauma, skeletal pathology, or skeletal anomalies. Other individualizing traits include the shape of various bones or their features, such as the frontal sinuses (Figure 15.16).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Comparative dental radiography focuses on the number, shape, location, and orientation of dentition and dental restorations in antemortem and postmortem images. While there is not a minimum number of matching traits that need to be identified for an identification to be made, the antemortem and postmortem records should have enough skeletal or dental consistencies to conclude that the records did in fact come from the same individual (SWGANTH 2010a). Consideration should also be given to population-level frequencies of specific skeletal and dental traits. If a trait is particularly common within a given population, it may not be a good trait to utilize for positive identification.<\/p>\n<figure style=\"width: 354px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image16-3.png\" alt=\"A scapula and humerus with a metal shoulder replacement.\" width=\"354\" height=\"231\" \/><figcaption class=\"wp-caption-text\">Figure 15.17: Image of joint replacement in the right shoulder. Credit: <a href=\"https:\/\/naturalhistory.si.edu\/education\/teaching-resources\/written-bone\/skeleton-keys\/todays-bones\">Shoulder replacement<\/a> by <a href=\"https:\/\/www.si.edu\/\">Smithsonian<\/a> [exhibit: Written in Bone, Today\u2019s Bones] <a href=\"https:\/\/www.si.edu\/termsofuse\">is used for educational and non-commercial purposes as outlined by the Smithsonian.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Surgical implants or devices can also be used for identification purposes (Figure 15.17). These implements are sometimes recovered with human remains. One of the ways forensic anthropologists can use surgical implants to assist in decedent identification is by providing a thorough analysis of the implant and noting any identifying information such as serial numbers, manufacturer symbols, and so forth. This information can then sometimes be tracked directly to the manufacturer or the place of surgical intervention, which may be used to identify unknown remains (SWGANTH 2010a).<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Trans Doe Task Force<\/h2>\n<p class=\"import-Normal\">The Trans Doe Task Force (TDTF) is a Trans-led nonprofit organization that investigates cases involving LGBTQ+ missing and murdered persons. The organization specifically focuses on transgender and gender-variant cases, providing connections between law enforcement agencies, medical examiner offices, forensic anthropologists, and forensic genetic genealogists to increase the chances of identification. Additionally, the TDTF curates a data repository of missing, murdered, and unclaimed LGBTQ+ individuals, and they continuously try innovative approaches to identify these individuals, whose lived gender identity may not match their biological sex.<\/p>\n<p class=\"import-Normal\">For more information visit <a href=\"https:\/\/transdoetaskforce.org\/\">transdoetaskforce.org<\/a><\/p>\n<\/div>\n<h3 class=\"import-Normal\"><strong>Trauma Analysis<\/strong><\/h3>\n<h4 class=\"import-Normal\"><em>Types of Trauma<\/em><strong><br \/>\n<\/strong><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Within the field of anthropology, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1256\">trauma<\/a> <\/strong>is defined as an injury to living tissue caused by an extrinsic force or mechanism (Lovell 1997:139). Forensic anthropologists can assist a forensic pathologist by providing an interpretation of the course of events that led to skeletal trauma. Typically, traumatic injury to bone is classified into one of four categories, defined by the trauma mechanism. A trauma mechanism refers to the force that produced the skeletal modification and can be classified as (1) sharp force, (2) blunt force, (3) projectile, or (4) thermal (burning). Each type of trauma, and the characteristic pattern(s) associated with that particular categorization, will be discussed below.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">First, let\u2019s consider s<em>harp-force trauma<\/em>, which is caused by a tool that is edged, pointed, or beveled\u2014for example, a knife, saw, or machete (SWGANTH 2011). The patterns of injury resulting from sharp-force trauma include linear incisions created by a sharp, straight edge; punctures; and chop marks (Figure 15.18; SWGANTH 2011). When observed under a microscope, an anthropologist can often determine what kind of tool created the bone trauma. For example, a power saw cut will be discernible from a manual saw cut.<\/p>\n<figure style=\"width: 602px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image21-1.png\" alt=\"Anterior image of a skull with multiple traumatic injuries to forehead.\" width=\"602\" height=\"457\" \/><figcaption class=\"wp-caption-text\">Figure 15.18: Example of sharp-force trauma (sword wound) to the frontal bone. The skull appears sliced with thin lines in two places across the top of the skull. Credit: <a href=\"https:\/\/openverse.org\/image\/909d1b77-ad5f-4cda-be44-6d9b5fbf14b9\/\">Female skull injured by a medieval sword<\/a> by <a href=\"https:\/\/sketchfab.com\/provinciaal_depot_noordholland\">Provinciaal depot voor archeologie Noord-Holland<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY 4.0 License<\/a>. The original image is a 3D model that can be manipulated on the <a href=\"https:\/\/wordpress.org\/openverse\/image\/909d1b77-ad5f-4cda-be44-6d9b5fbf14b9\/\">openverse website<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Second, <em>blunt-force trauma<\/em> is defined as \u201ca relatively low-velocity impact over a relatively large surface area\u201d (Galloway 1999, 5). Blunt-force injuries can result from impacts from clubs, sticks, fists, and so forth. Blunt-force impacts typically leave an injury at the point of impact but can also lead to bending and deformation in other regions of the bone. Depressions, fractures, and deformation at and around the site of impact are all characteristics of blunt-force trauma (Figure 15.19). As with sharp-force trauma, an anthropologist attempts to interpret blunt-force injuries, providing information pertaining to the type of tool used, the direction of impact, the sequence of impacts, if more than one, and the amount of force applied.<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 578px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image30.png\" alt=\"Cranium with two blunt force impacts from a hammer.\" width=\"578\" height=\"803\" \/><figcaption class=\"wp-caption-text\">Figure 15.19: Example of multiple blunt force impacts to the left parietal and frontal bones. There is one hole in the skull with fractured bone around the edges. There are also multiple spots across the back of the skull with depressions of various sizes. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Skull_hammer_trauma.jpg\">Skull hammer trauma<\/a> by <a href=\"https:\/\/www.nih.gov\/\">the National Institutes of Health<\/a>, Health &amp; Human Services, is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>. [Exhibit: Visible Proofs: Forensic Views of the Body, U.S. National Library of Medicine, 19th Century Collection, National Museum of Health and Medicine, Armed Forces Institute of Pathology, Washington, D.C.]<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Third, <em>projectile trauma<\/em> refers to high-velocity trauma, typically affecting a small surface area (Galloway 1999, 6). Projectile trauma results from fast-moving objects such as bullets or shrapnel. It is typically characterized by penetrating defects or embedded materials (Figure 15.20). When interpreting injuries resulting from projectile trauma, an anthropologist can often offer information pertaining to the type of weapon used (e.g., rifle vs. handgun), relative size of the bullet (but not the caliber of the bullet), the direction the projectile was traveling, and the sequence of injuries if there are multiple present.<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 462px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image5-3.png\" alt=\"Anterior and posterior views of a skull with a gunshot wound.\" width=\"462\" height=\"291\" \/><figcaption class=\"wp-caption-text\">Figure 15.20: Example of projectile trauma with an entrance wound to the frontal bone and exit wound visible on the occipital. A small circular hole is visible in the front of the skull with cracks radiating out from the point of impact. There is a larger hole visible in the back of the skull that is irregular yet circular in shape. Credit: <a href=\"https:\/\/naturalhistory.si.edu\/education\/teaching-resources\/written-bone\/skeleton-keys\/how-bone-biographies-get-written\">Trauma: Gunshot Wounds<\/a> by <a href=\"https:\/\/www.si.edu\/\">Smithsonian<\/a> [exhibit: Written in Bone, How Bone Biographies Get Written] <a href=\"https:\/\/www.si.edu\/termsofuse\">is used for educational and non-commercial purposes as outlined by the Smithsonian.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Finally, <em>thermal trauma<\/em> is a bone alteration that results from bone exposure to extreme heat. Thermal trauma can result in cases of house or car fires, intentional disposal of a body in cases of homicidal violence, plane crashes, and so on. Thermal trauma is most often characterized by color changes to bone, ranging from yellow to black (charred) or white (calcined). Other bone alterations characteristic of thermal trauma include delamination (flaking or layering due to bone failure), shrinkage, fractures, and heat-specific burn patterning. When interpreting injuries resulting from thermal damage, an anthropologist can differentiate between thermal fractures and fractures that occurred before heat exposure, thereby contributing to the interpretation of burn patterning (e.g., was the individual bound or in a flexed position prior to the fire?).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">While there are characteristic patterns associated with the four categories of bone trauma, it is also important to note that these bone alterations do not always occur independently of different trauma types. An individual\u2019s skeleton may present with multiple different types of trauma, such as a projectile wound and thermal trauma. Therefore, it is important that the anthropologist recognize the different types of trauma and interpret them appropriately.<\/p>\n<h3 class=\"import-Normal\"><strong>Timing of Injury<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Another important component of any anthropological trauma analysis is the determination of the timing of injury (e.g., when did the injury occur). Timing of injury is traditionally split into one of three categories: <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1260\">antemortem<\/a> <\/strong>(before death), <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1258\">perimortem<\/a> <\/strong>(at or around the time of death), and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1266\">postmortem <\/a><\/strong>(after death). This classification system differs slightly from the classification system used by the pathologist because it specifically references the qualities of bone tissue and bone response to external forces. Therefore, the perimortem interval (at or around the time of death) means that the bone is still fresh and has what is referred to as a green bone response, which can extend past death by several weeks or even months. For example, in cold or freezing temperatures a body can be preserved for extended periods of time, increasing the perimortem interval, while in desert climates decomposition is accelerated, thereby significantly decreasing the postmortem interval (Galloway 1999, 12). Antemortem injuries (occurring well before death and not related to the death incident) are typically characterized by some level of healing, in the form of a fracture callus or unification of fracture margins. Finally, postmortem injuries (occurring after death, while bone is no longer fresh) are characterized by jagged fracture margins, resulting from a loss of moisture content during the decomposition process (Galloway 1999, 16). In general, all bone traumas should be classified according to the timing of injury, if possible. This information will help the medical examiner or pathologist better understand the circumstances surrounding the decedent\u2019s death, as well as events occurring during life and after the final disposition of the body.<\/p>\n<h3 class=\"import-Normal\"><strong>The Role of the Forensic Anthropologist in Trauma Analysis<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Within the medicolegal system, forensic anthropologists are often called upon by the medical examiner, forensic pathologist, or coroner to assist with an interpretation of trauma. The forensic anthropologist\u2019s main focus in any trauma analysis is the underlying skeletal system\u2014as well as, sometimes, cartilage. Analysis and interpretation of soft tissue injuries fall within the purview of the medical examiner or pathologist. It is also important to note that the main role of the forensic anthropologist is to provide information pertaining to skeletal injury to assist the medical examiner\/pathologist in their final interpretation of injury. Forensic anthropologists do not hypothesize as to the cause of death of an individual. Instead, a forensic anthropologist\u2019s report should include a description of the injury (e.g., trauma mechanism, number of injuries, location, timing of injury); documentation of the injury, which may be utilized in court testimony (e.g., photographs, radiographs, measurements); and, if applicable, a statement as to the condition of the body and state of decomposition, which may be useful for understanding the depositional context (e.g., how long has the body been exposed to the elements; was it moved or in its original location; are any of the alterations to bone due to environmental or faunal exposure instead of intentional human modification).<\/p>\n<h2 class=\"import-Normal\">Taphonomy<\/h2>\n<h2 class=\"import-Normal\"><strong>What Happened to the Remains After Death?<\/strong><\/h2>\n<p class=\"import-Normal\">The majority of the skeletal analysis process revolves around the identity of the deceased individual. However, there is one last, very important question that forensic anthropologists should ask: What happened to the remains after death? Generally speaking, processes that alter the bone after death are referred to as taphonomic changes (refer to Chapter 7 for a discussion regarding taphonomy and the fossil record).<\/p>\n<p class=\"import-Normal\">The term <em>taphonomy<\/em> was originally used to refer to the processes through which organic remains mineralize, also known as fossilization. Within the context of biological anthropology, the term <em>taphonomy<\/em> is better defined as the study of what happens to human remains after death (Komar and Buikstra 2008). Initial factors affecting a body after death include processes such as decomposition and scavenging by animals. However, taphonomic processes encompass much more than the initial period after death. For example, plant root growth can leach minerals from bone, leaving a distinctive mark. Sunlight can bleach human remains, leaving exposed areas whiter than those that remained buried. Water can wear the surface of the bone until it becomes smooth.<\/p>\n<p class=\"import-Normal\">Some taphonomic processes can help a forensic anthropologist estimate the relative amount of time that human remains have been exposed to the elements. For example, root growth through a bone would certainly indicate a body was buried for more than a few days. Forensic anthropologists must be very careful when attempting to estimate time since death based on taphonomic processes because environmental conditions can greatly influence the rate at which taphonomic processes progress. For example, in cold environments, tissue may decay slower than in warm, moist environments.<\/p>\n<p class=\"import-Normal\">Forensic anthropologists must contend with taphonomic processes that affect the preservation of bones. For example, high acidity in the soil can break down human bone to the point of crumbling. In addition, when noting trauma, they must be very careful not to confuse postmortem (after death) bone damage with trauma.<\/p>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 470.25pt\">\n<caption>Figure 15.21: Table showing taphonomic processes that affect the preservation of bones. A. Rodent gnawing. B. Carnivore damage. C. Burned bone. D. Root etching. E. Weathering. F. Cut marks. Credit: A. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Rodent gnawing (Figure 15.26)<\/a>, B. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Carnivore damage (Figure 15.27)<\/a>, C. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Burned bone (Figure 15.28)<\/a>, D. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Root etching (Figure 15.29)<\/a>, E. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Weathering (Figure 15.30)<\/a>, and F. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Cut marks (Figure 15.30)<\/a>, all original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone are under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-NC 4.0 License<\/a>.<\/caption>\n<thead>\n<tr style=\"height: 52.5pt\">\n<td class=\"Table1-C\" style=\"padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Taphonomic Process<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 1pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Definition<\/p>\n<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table1-R\" style=\"height: 190.5pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Rodent Gnawing<\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: center\"><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image19-2.png\" alt=\"Parallel tooth marks etched by a rodent\u2019s front teeth visible on the end of an animal bone.\" width=\"564\" height=\"422\" \/><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">When rodents, such as rats and mice, chew on bone, they leave sets of parallel grooves. The shallow grooves are etched by the rodent\u2019s incisors.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 166.75pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Carnivore Damage<\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: center\"><strong><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image23-4.png\" alt=\"Pit marks from the canines of a carnivore visible on the surface of an animal bone.\" width=\"410\" height=\"272\" \/><\/strong><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Carnivores may leave destructive dental marks on bone. The tooth marks may be visible as pit marks or punctures from the canines, as well as extensive gnawing or chewing of the ends of the bones to retrieve marrow.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 177pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Burned Bone<\/strong><\/p>\n<p class=\"import-Normal\"><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image20-5.png\" alt=\"Burned animal bone fragments pictured at different stages of thermal damage.\" width=\"512\" height=\"342\" \/><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Fire causes observable damage to bone. Temperature and the amount of time bone is heated affect the appearance of the bone. Very high temperatures can crack bone and result in white coloration. Color gradients are visible in between high and lower temperatures, with lower temperatures resulting in black coloration from charring. Cracking can also reveal information about the directionality of the burn.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 169.75pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Root Etching<\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: center\"><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image15-4.png\" alt=\"Animal bone with prominent, discolored grooves where roots leached nutrients from bone\u2019s surface.\" width=\"512\" height=\"342\" \/><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Plant roots can etch the outer surface of bone, leaving grooves where the roots attached as they leached nutrients. During this process, the plant\u2019s roots secrete acid that breaks down the surface of the bone.<\/p>\n<p class=\"import-Normal\">\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 170.5pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Weathering<\/strong><\/p>\n<p class=\"import-Normal\"><strong><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image9.png\" alt=\"Cracking and exfoliation of the surface of an animal bone. \" width=\"512\" height=\"342\" \/><\/strong><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Many different environmental conditions affect bone. River transport can smooth the surface of the bone due to water abrasion. Sunlight can bleach the exposed surface of bone. Dry and wet environments or the mixture of both types of environments can cause cracking and exfoliation of the surface. Burial in different types of soil can cause discoloration, and exposure can cause degreasing.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 169.75pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Cut Marks<\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: left\"><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image8-2.png\" alt=\"Thin vertical lines and cuts are visible along the bone.\" width=\"512\" height=\"342\" \/><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Humans may alter bone by cutting, scraping, or sawing it directly or in the process of removing tissue. The groove pattern\u2014that is, the depth and width of the cuts\u2014can help identify the tool used in the cutting process.<\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div class=\"textbox textbox--sidebar shaded\">\n<p>Type your textbox content here.<\/p>\n<\/div>\n<p>&nbsp;<\/p>\n<h2>Dig Deeper: Modern Forensic Technologies<\/h2>\n<p>In recent years, the forensics community has greatly benefited from the introduction of new technologies, helping strengthen the precision and speed of discoveries and advancements in the field. With recent developments in forensic anthropology, such as 3D scanning technologies, virtual reconstruction, and AI-assisted DNA analysis being integrated into traditional methods, there have been notable changes in how experts investigate human remains.<\/p>\n<p><strong>Artificial intelligence<\/strong><\/p>\n<p>In recent years, Artificial intelligence (AI) has shown itself to be a valuable tool within forensic anthropology. Aiding forensic experts and toxicologists with complex tasks, the limitations of traditional autopsies can be addressed with the help of AI. By automating and enhancing key investigative processes such as searching for microscopic changes in the human body to determine the cause of death or a person\u2019s life conditions, AI has the potential to enhance the efficiency of forensic processes significantly. It facilitates the detection of microscopic bodily changes to determine the cause of death or living conditions, compares evidence against databases for weapon identification and blood spatter analysis, and reduces manual workload. AI also enables the electronic storage of biometric data\u2013such as facial features, retinal patterns, and fingerprints\u2013for more accurate identity verification. Additionally, AI-powered microscopy enhances the detection of biological traces on complex surfaces, while blood biomarker analysis allows for more precise estimations of time of death (Wankhade et al., 2022).<\/p>\n<p>While AI holds great promise for the future of forensic medicine, a significant challenge remains: sourcing high-quality data to train the algorithms effectively. One of the more recent AI technologies making waves in the forensic anthropology sector is a new automated AI algorithm called the Convolutional Neural Network (CNN). As described by researchers in Switzerland\u2019s national medical journal Healthcare, CNN is a Deep Learning algorithm that allows for the detection of microscopic skull damage from CT scans or soft-tissue predictions of a face based on the skull information provided (Thurzo et al., 2021). While there are many advantages to using the CNN, the algorithm can be subject to biases in the same way human forensic experts can, as its assessment and pattern recognition of skulls and skeletons depend on the source data initially used for its AI training (2021).<\/p>\n<p><strong>3D Modeling<\/strong><\/p>\n<p>Identifying complex trauma to bones\u2013such as distinguishing heat fractures following blunt force trauma\u2013remains a significant challenge in forensic anthropology. This is particularly true for irregular skeletal structures like the pelvis, where overlapping trauma types can be difficult to differentiate, leading to these bones often being understudied. A 2024 study done by researchers from the University of Alberta in collaboration with the Michigan State Police explores the use of 3D laser scans and modelling technology to provide a highly detailed analysis of irregular bones with trauma. The study aimed to better distinguish peri-mortem trauma (trauma occurring around the time of death) from post-mortem heat alterations and improve the forensic analysis accuracy of such cases (Friedlander et al., 2024). The use of 3D laser scans and modelling technology provides very clear, detailed, and colored scans of bones, showing distinctions between the characteristics of the fractures. Blunt force and sharp force trauma produce a colour gradient on the 3D model that is more gradual and irregular, while heat fractures are more neat and characterized by little colour variation on the 3D models (2024). Other conclusions were also drawn from the study, such as the differences in trauma on fresh bones and bones that have been exposed to the elements for longer. An example of this is the interstitial fluid and collagen fibrils in fresh bones absorbing force, causing more long and jagged fracture lines, as opposed to a brittle fracture that older bones may exhibit (2024).<\/p>\n<p>Overall, the integration of 3D modeling technology offers a reproducible and highly detailed approach for analyzing trauma in anatomically complex and historically understudied skeletal regions. The practicality of this advancement is further emphasized by the researchers, who note that \u201cin many instances, scanned 3D models can be 3D printed for handheld representation of the model without damaging or overhandling the remains\u201d (2024, p. 2). By enhancing the ability to differentiate between various types of trauma and allowing for more convenient and risk-averse methods of research, this technology significantly improves the accuracy and reliability of forensic interpretations.<\/p>\n<h2>Ethics and Human Rights<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Working with human remains requires a great deal of consideration and respect for the dead. Forensic anthropologists have to think about the ethics of our use of human remains for scientific purposes. How do we conduct casework in the most respectable manner possible? While there are a wide range of ethical considerations to consider when contemplating a career in forensic anthropology, this chapter will focus on two major categories: working with human remains and acting as an expert within the medicolegal system.<\/p>\n<h3 class=\"import-Normal\"><strong>Working with Human Remains<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropologists work with human remains in a number of contexts, including casework, excavation, research, and teaching. When working with human remains, it is always important to use proper handling techniques. To prevent damage to skeletal remains, bones should be handled over padded surfaces. Skulls should never be picked up by placing fingers in the eye orbits, foramen magnum (hole at the base of the skull for entry of the spinal cord), or through the zygomatic arches (cheekbones). Human remains, whether related to casework, fieldwork, donated skeletal collections, or research, were once living human beings. It is important to always bear in mind that work with remains should be ingrained with respect for the individual and their relatives. In addition to fieldwork, casework, and teaching, anthropologists are often invited to work with remains that come from a bioarchaeological context or from a human rights violation. While this discussion of ethics is not comprehensive, two case examples will be provided below in which an anthropologist must consider the ethical standards outlined above.<\/p>\n<h3 class=\"import-Normal\"><strong>Modern Human Rights Violations<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropologists may also be called to participate in criminal investigations involving human rights violations. Anthropological investigations may include assistance with identifications, determination of the number of victims, and trauma analyses. In this role, forensic anthropologists play an integral part in promoting human rights, preventing future human rights violations, and providing the evidence necessary to prosecute those responsible for past events. A few ethical considerations for the forensic anthropologist involved in human rights violations include the use of appropriate standards of identification, presenting reliable and unbiased testimony, and maintaining preservation of evidence. For a more comprehensive history of forensic anthropological contributions to human rights violations investigations, see Ubelaker 2018.<\/p>\n<h3 class=\"import-Normal\"><strong>Acting as an Expert in the Medicolegal System<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In addition to the ethical considerations involved in working with human skeletal remains, forensic anthropologists must abide by ethical standards when they act as experts within the medicolegal system. The role of the forensic anthropologist within the medicolegal system is primarily to provide information to the medical examiner or coroner that will aid in the identification process or determination of cause and manner of death. Forensic anthropologists also may be called to testify in a court of law. In this capacity, forensic anthropologists should always abide by a series of ethical guidelines that pertain to their interpretation, presentation, and preservation of evidence used in criminal investigations. First and foremost, practitioners should never misrepresent their training or education. When appropriate, outside opinions and assistance in casework should be requested (e.g., consulting a radiologist for radiological examinations or odontologist for dental exams). The best interest of the decedent should always take precedence. All casework should be conducted in an unbiased way, and financial compensation should never be accepted as it can act as an incentive to take a biased stance regarding casework. All anthropological findings should be kept confidential, and release of information is best done by the medical examiner or coroner. Finally, while upholding personal ethical standards, forensic anthropologists are also expected to report any perceived ethical violations committed by their peers.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ethical standards for the field of forensic anthropology are outlined by the Organization of Scientific Area Committees (OSAC) for Forensic Science, administered by the National Institute of Standards and Technology (NIST). OSAC and NIST recently began an initiative to develop standards that would strengthen the practice of forensic science both in the United States and internationally. OSAC\u2019s main objective is to \u201cstrengthen the nation\u2019s use of forensic science by facilitating the development of technically sound forensic science standards and by promoting the adoption of those standards by the forensic science community\u201d (NIST n.d.). Additionally, OSAC promotes the establishment of best practices and other guidelines to ensure that forensic science findings and their presentation are reliable and reproducible (NIST 2023).<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Native American Graves Protection and Repatriation Act (NAGPRA)<\/h2>\n<p class=\"import-Normal\">There is a long history in the <span style=\"background-color: #00ffff\">United States<\/span> of systematic disenfranchisement of Native American people, including lack of respect for tribal sovereignty. This includes the egregious treatment of Native American human remains. Over several centuries, thousands of Native American remains were removed from tribal lands and held at institutions in the United States, such as museums and universities.<\/p>\n<p class=\"import-Normal\">In 1990, a landmark human rights federal law, the Native American Graves Protection and Repatriation Act (NAGPRA), spurred change in the professional standards and practice of biological anthropology and archaeology. NAGPRA established a legal avenue to provide protection for and repatriation of Native American remains, cultural items, and sacred objects removed from Federal or tribal lands to Native American lineal descendants, Indian tribes, and Native Hawaiian organizations. Human remains and associated artifacts, curated in museum collections and federally funded institutions, are subject to three primary provisions outlined by the NAGPRA statute: (1) protection for Native graves on federal and private land; (2) recognition of tribal authority on such lands; and (3) the requirement that all Native skeletal remains and associated artifacts be inventoried and culturally affiliated groups be consulted concerning decisions related to ownership and final disposition (Rose, Green, and Green 1996). NAGPRA legislation was enacted to ensure ethical consideration and treatment of Native remains and to improve dialogue between scientists and Native groups.<\/p>\n<ul>\n<li>For more information about NAGPRA, visit the <a href=\"https:\/\/www.usbr.gov\/nagpra\/\" target=\"_blank\" rel=\"noopener\">Bureau of Reclamation NAGPRA website<\/a><\/li>\n<li>To read the text of the law, visit the <a href=\"https:\/\/www.congress.gov\/bill\/101st-congress\/house-bill\/5237\">US Congress NAGPRA law website<\/a>.<\/li>\n<li>For further discussion of NAGPRA history, please see <a href=\"https:\/\/textbooks.whatcom.edu\/tracesarchaeology\/\" target=\"_blank\" rel=\"noopener\"><em>TRACES: <\/em><em>An Open Invitation to <\/em><em>Archaeology <\/em>open textbook website<\/a><em><br \/>\n<\/em><\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Becoming a Forensic Anthropologist<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"background-color: #ff99cc\">What does it take to be a forensic anthropologist? Forensic anthropologists are first and foremost anthropologists. While many forensic anthropologists have an undergraduate degree in anthropology, they may also major in biology, criminal justice, pre-law, pre-med, and many other related fields. Practicing forensic anthropologists typically have an advanced degree, either a Master\u2019s or Doctoral degree in Anthropology. Additional training and experience in archaeology, the medico-legal system, rules of evidence, and expert witness testimony are also common. Practicing forensic anthropologists are also encouraged to be board-certified through the American Board of Forensic Anthropology (ABFA). Learn more about the field and educational opportunities on the ABFA website: <a class=\"rId111\" style=\"background-color: #ff99cc\" href=\"https:\/\/www.theabfa.org\/coursework\">https:\/\/www.theabfa.org\/coursework<\/a>.<\/span><\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Review Questions<\/h2>\n<ul>\n<li>What is forensic anthropology? What are the seven primary steps involved in a skeletal analysis?<\/li>\n<li>What are the major components of a biological profile? Why are forensic anthropologists often-tasked with creating biological profiles for unknown individuals?<\/li>\n<li>What are the four major types of skeletal trauma?<\/li>\n<li>What is taphonomy, and why is an understanding of taphonomy often critical in forensic anthropology analyses?<\/li>\n<li>What are some of the ethical considerations faced by forensic anthropologists?<\/li>\n<\/ul>\n<\/div>\n<h2>About the Authors<\/h2>\n<p><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image24-4.jpg\" alt=\"A woman with straight blonde hair smiles at the camera. \" width=\"191\" height=\"254\" \/><\/p>\n<h3 class=\"import-Normal\"><strong>Ashley Kendell, Ph.D.<\/strong><\/h3>\n<p class=\"import-Normal\">California State University, Chico, <a class=\"rId113\" href=\"mailto:akendell@csuchico.edu\">akendell@csuchico.edu<\/a><\/p>\n<p class=\"import-Normal\">Dr. Ashley Kendell is currently an associate professor and forensic anthropologist at Chico State. Prior to beginning her position at Chico State, she was a visiting professor at the University of Montana and the forensic anthropologist for the state of Montana. Dr. Kendell obtained her doctorate from Michigan State University, and her research interests include skeletal trauma analysis and digitization and curation methods for digital osteological data. She is also a Registry Diplomate of the American Board of Medicolegal Death Investigators. Throughout her doctoral program, she worked as a medicolegal death investigator for the greater Lansing, Michigan, area and was involved in the investigation of over 200 forensic cases.<\/p>\n<p class=\"import-Normal\"><strong><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image4.jpg\" alt=\"A woman with straight brown hair pulled back smiles at the camera. \" width=\"194\" height=\"258\" \/><\/strong><\/p>\n<h3 class=\"import-Normal\"><strong>Alex Perrone, M.A., M.S.N, R.N., P.H.N.<\/strong><\/h3>\n<p class=\"import-Normal\">Butte Community College, <a class=\"rId115\" href=\"mailto:perroneal@butte.edu\">perroneal@butte.edu<\/a><\/p>\n<p class=\"import-Normal\">Alex Perrone is a lecturer in anthropology at Butte Community College. She is also a Registered Nurse and a certified Public Health Nurse. She is a former Supervisor of the Human Identification Laboratory in the Department of Anthropology at California State University, Chico. Her research interests include bioarchaeology, paleopathology, forensic anthropology, skeletal biology, California prehistory, and public health. She has worked on bioarchaeological and archaeological projects in Antigua, California, Hawaii, Greece, and the UK, and was an archaeological technician for the USDA Forest Service. She assisted with training courses for local and federal law enforcement agencies and assisted law enforcement agencies with the recovery and analysis of human remains.<\/p>\n<p class=\"import-Normal\" data-wp-editing=\"1\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image1-1.jpg\" alt=\"A woman with curly brown, shoulder-length hair smiles at the camera.\" width=\"190\" height=\"253\" \/><\/p>\n<h3 class=\"import-Normal\"><strong>Colleen Milligan, Ph.D.<\/strong><\/h3>\n<p class=\"import-Normal\">California State University, Chico, <a class=\"rId117\" href=\"mailto:cfmilligan@csuchico.edu\">cfmilligan@csuchico.edu<\/a><\/p>\n<p class=\"import-Normal\">Dr. Colleen Milligan is a biological and forensic anthropologist with research interests in bioarchaeology, skeletal biology, and forensic anthropology. She has been a Fellow with the Department of Homeland Security and has assisted in forensic anthropology casework and recoveries in the State of Michigan and California. She has also assisted in community outreach programs in forensic anthropology and forensic science, as well as recovery training courses for local, state, and federal law enforcement officers. She is a certified instructor through Peace Officers Standards and Training (POST). Dr. Milligan serves as the current co-director of the Chico State Human Identification Laboratory.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration<\/h2>\n<p><a href=\"https:\/\/www.theabfa.org\/coursework\" target=\"_blank\" rel=\"noopener\">The American Board of Forensic Anthropology (ABFA)<\/a><\/p>\n<p><a href=\"https:\/\/www.aafs.org\/\" target=\"_blank\" rel=\"noopener\">The American Academy of Forensic Sciences (AAFS)<\/a><\/p>\n<p><a href=\"https:\/\/www.nist.gov\/organization-scientific-area-committees-forensic-science\" target=\"_blank\" rel=\"noopener\">The Organization of Scientific Area Committees for Forensic Science (OSAC)<\/a><\/p>\n<p><a href=\"https:\/\/textbooks.whatcom.edu\/tracesarchaeology\/\" target=\"_blank\" rel=\"noopener\">TRACES Bioarchaeology<\/a><\/p>\n<p><a href=\"https:\/\/transdoetaskforce.org\/\" target=\"_blank\" rel=\"noopener\">Trans Doe Task Force<\/a><\/p>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Adams, Bradley J., and Lyle W. Konigsberg, eds. 2008. <em>Recovery, Analysis, and Identification of Commingled Remains<\/em>. Totowa, NJ: Humana Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Beatrice, Jared S., and Angela Soler. 2016. \u201cSkeletal Indicators of Stress: A Component of the Biocultural Profile of Undocumented Migrants in Southern Arizona.\u201d <em>Journal of Forensic Sciences <\/em>61 (5): 1164\u20131172.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Berg, Gregory E. 2017. \u201cSex Estimation of Unknown Human Skeletal Remains.\u201d In <em>Forensic Anthropology: A Comprehensive Introduction, Second Edition<\/em>, edited by Natalie R. Langley and MariaTeresa A. Tersigni-Tarrant, 143\u2013159. Boca Raton, FL: CRC Press.<\/p>\n<p class=\"import-Normal\">Bethard, Jonathan D., and Elizabeth A. DiGangi. 2020. \u201cLetter to the Editor\u2014Moving Beyond a Lost Cause: Forensic Anthropology and Ancestry Estimates in the United States.\u201d <em>Journal of Forensic Sciences<\/em> 65 (5): 1791\u20131792.<\/p>\n<p class=\"import-Normal\">Birkby, Walter H., Todd W. Fenton, and Bruce E. Anderson. 2008. \u201cIdentifying Southwest Hispanics Using Nonmetric Traits and the Cultural Profile.\u201d <em>Journal of Forensic Sciences <\/em>53 (1): 29\u201333.<\/p>\n<p class=\"import-Normal\">Blatt, Samantha, Amy Michael, and Lisa Bright. Forthcoming. \u201cBioarchaeology: Interpreting Human Behavior from Skeletal Remains.\u201d In <em>TRACES: <\/em><em>An Open Invitation to <\/em><em>Archaeology<\/em>. https:\/\/textbooks.whatcom.edu\/tracesarchaeology\/.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Brooks, S., and J. M. Suchey. 1990. \u201cSkeletal Age Determination Based on the Os Pubis: A Comparison of the Acs\u00e1di-Nemesk\u00e9ri and Suchey-Brooks Methods.\u201d <em>Human Evolution <\/em>5 (3): 227\u2013238.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Buchanan, Shelby. 2014. \u201cBone Modification in Male to Female Transgender Surgeries: Considerations for the Forensic Anthropologist.\u201d MA thesis, Department of Geography and Anthropology, Louisiana State University, Baton Rouge.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Cunningham, Craig, Louise Scheuer, and Sue Black. 2016. <em>Developmental Juvenile Osteology, Second Edition<\/em>. London: Elsevier Academic Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Galloway, Alison, ed. 1999. <em>Broken Bones: Anthropological Analysis of Blunt Force Trauma<\/em>. Springfield, IL: Charles C. Thomas Publisher, LTD.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Hefner, Joseph T., and Kandus C. Linde. 2018. <em>Atlas of Human Cranial <\/em><em>Macromorphoscopic<\/em><em> Traits<\/em>. San Diego: Academic Press.<\/p>\n<p class=\"import-Normal\">\u0130\u015fcan, M. Y., S. R. Loth, and R. K. Wright. 1984. \u201cAge Estimation from the Rib by Phase Analysis: White Males.\u201d <em>Journal of Forensic Sciences <\/em>29 (4): 1094\u20131104.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">\u0130\u015fcan, M. Y., S. R. Loth, and R. K. Wright. 1985. \u201cAge Estimation from the Rib by Phase Analysis: White Females.\u201d <em>Journal of Forensic Sciences <\/em>30 (3): 853\u2013863.Katz, Darryl, and Judy Myers Suchey. 1986. \u201cAge Determination of the Male Os Pubis.\u201d <em>American Journal of Physical Anthropology <\/em>69 (4): 427\u2013435.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Komar, Debra A., and Jane E. Buikstra. 2008. <em>Forensic Anthropology: Contemporary Theory and Practice<\/em>. New York: Oxford University Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Langley, Natalie R., Alice F. Gooding, and MariaTeresa Tersigni-Tarrant. 2017. \u201cAge Estimation Methods.\u201d In <em>Forensic Anthropology: A Comprehensive Introduction, Second Edition<\/em>, edited by Natalie R. Langley and MariaTeresa A. Tersigni-Tarrant, 175\u2013191. Boca Raton, FL: CRC Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Lovell, Nancy C. 1997. \u201cTrauma Analysis in Paleopathology.\u201d <em>Yearbook of Physical Anthropology<\/em> 104 (S25): 139\u2013170.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Native American Graves Protection and Repatriation Act (NAGPRA) 1990 (25 U.S. Code 3001 et seq.)<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">NIST (National Institute of Standards and Technology). N.d. \u201cThe Organization of Scientific Area Committees for Forensic Science.\u201d Accessed April 18, 2023. <a class=\"rId120\" href=\"https:\/\/www.nist.gov\/topics\/organization-scientific-area-committees-forensic-science\">https:\/\/www.nist.gov\/topics\/organization-scientific-area-committees-forensic-science<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ousley, Stephen. 1995. \u201cShould We Estimate Biological or Forensic Stature?\u201d <em>Journal of Forensic Sciences<\/em> 40(5): 768\u2013773.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Phenice, T. W. 1969. \u201cA Newly Developed Visual Method of Sexing the Os Pubis.\u201d <em>American Journal of Physical Anthropology<\/em> 30 (2): 297\u2013302.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Rose, Jerome C., Thomas J. Green, and Victoria D. Green. 1996. \u201cNAGPRA Is Forever: Osteology and the Repatriation of Skeletons.\u201d <em>Annual Review of Anthropology <\/em>25: 81\u2013103.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Schaefer, Maureen, Sue Black, and Louise Scheuer. <em>Juvenile Osteology: A Laboratory and Field Manua<\/em>l. 2009. San Diego: Elsevier Academic Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Schall, Jenna L., Tracy L. Rogers, and Jordan D. Deschamps-Braly. 2020. \u201cBreaking the Binary: The Identification of Trans-women in Forensic Anthropology.\u201d <em>Forensic Science International<\/em> 309: 110220. https:\/\/doi.org\/10.1016\/j.forsciint.2020.110220.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2010a. \u201cPersonal Identification.\u201d Last modified June 3, 2010. <a class=\"rId121\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_personal_identification.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_personal_identification.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2010b. \u201cSex Assessment.\u201d Last modified June 3, 2010. <a class=\"rId122\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_sex_assessment.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_sex_assessment.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2011. \u201cTrauma Analysis.\u201d Last modified May 27, 2011. <a class=\"rId123\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_trauma.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_trauma.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2012. \u201cStature Estimation.\u201d Last modified August 2, 2012. <a class=\"rId124\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_stature_estimation.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_stature_estimation.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2013. \u201cAge Estimation.\u201d Last modified January 22, 2013. <a class=\"rId125\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_age_estimation.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_age_estimation.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Soler, Angela, and Jared S. Beatrice. 2018. \u201cExpanding the Role of Forensic Anthropology in Humanitarian Crisis: An Example from the USA-Mexico Border. In <em>Sociopolitics of Migrant Death and Repatriation: Perspectives from Forensic Science<\/em>, edited by Krista E. Latham and Alyson J. O\u2019Daniel, 115\u2013128. New York: Springer.<\/p>\n<p class=\"import-Normal\">Soler, Angela, Robin Reineke, Jared Beatrice, and Bruce E. Anderson. 2019. \u201cEtched in Bone: Embodied Suffering in the Remains of Undocumented Migrants.\u201d <em>In<\/em> <em>The Border and Its Bodies: The Embodiment of Risk along the U.S.-M\u00e9xico Line<\/em>, edited by Thomas E. Sheridan and Randall H. McGuire, 173\u2013207. Tucson: University of Arizona Press.<\/p>\n<p class=\"import-Normal\">Stull, Kyra E., Eric J. Bartelink, Alexandra R. Klales, Gregory E. Berg, Michael W. Kenyhercz, Erica N. L\u2019Abb\u00e9, Matthew C. Go, et al.. 2021. \u201cCommentary on: Bethard JD, DiGangi EA. Letter to the Editor\u2014Moving Beyond a Lost Cause: Forensic Anthropology and Ancestry Estimates in the United States. J Forensic Sci. 2020;65(5):1791\u20132. doi: 10.1111\/1556-4029.14513.\u201d <em>Journal of Forensic Sciences <\/em>66 (1): 417\u2013420.<\/p>\n<p class=\"import-Normal\">Tallman, Sean D., Caroline D. Kincer, and Eric D. Plemons. 2022. \u201cCentering Transgender Individuals in Forensic Anthropology and Expanding Binary Sex Estimation in Casework and Research.\u201d Special issue, \u201cDiversity and Inclusion,\u201d <em>Forensic Anthropology<\/em> 5 (2): 161\u2013180.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Tersigni-Tarrant, MariaTeresa A., and Natalie R. Langley. 2017. \u201cHuman Osteology.\u201d In <em>Forensic Anthropology: A Comprehensive Introduction, Second Edition<\/em>, edited by Natalie R. Langley and MariaTeresa A. Tersigni-Tarrant, 81\u2013109. Boca Raton, FL: CRC Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ubelaker, Douglas H. 2018. \u201cA History of Forensic Anthropology.\u201d Special issue, \u201cCentennial Anniversary Issue of AJPA,\u201d <em>American Journal of Physical Anthropology<\/em> 165 (4): 915\u2013923.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">White, Tim D., and Pieter A. Folkens. 2005. <em>The Human Bone Manual<\/em>. Burlington, MA: Elsevier Academic Press.<\/p>\n<p class=\"import-Normal\">Winburn, Allysha P., and Bridget Algee-Hewitt. 2021. \u201cEvaluating Population Affinity Estimates in Forensic Anthropology: Insights from the Forensic Anthropology Database for Assessing Methods Accuracy (FADAMA).\u201d <em>Journal of Forensic Sciences<\/em> 66 (4): 1210\u20131219.<\/p>\n<p class=\"import-Normal\">Winburn, Allysha Powanda, Sarah Kiley Schoff, and Michael W. Warren. 2016. \u201cAssemblages of the Dead: Interpreting the Biocultural and Taphonomic Signature of Afro- Cuban Palo Practice in Florida.\u201d <em>Journal of African Diaspora Archaeology and Heritage <\/em>5 (1): 1\u201337.<\/p>\n<\/div>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1782\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1782\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\">Ashley Kendell, Ph.D., California State University, Chico<\/p>\n<p class=\"import-Normal\">Alex Perrone, M.A., M.S.N, R.N., P.H.N., Butte Community College<\/p>\n<p class=\"import-Normal\">Colleen Milligan, Ph.D., California State University, Chico<\/p>\n<p class=\"import-Normal\"><em>This chapter is a revision from \"<\/em><a class=\"rId7\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\"><em>Chapter 15: Bioarchaeology and Forensic Anthropology<\/em><\/a><em>\u201d by Ashley Kendell, Alex Peronne, and Colleen Milligan. In <\/em><a class=\"rId8\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, first edition<\/em><\/a><em>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under <\/em><a class=\"rId9\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\"><em>CC BY-NC 4.0<\/em><\/a><em>. <\/em><\/p>\n<p class=\"import-Normal\"><strong>Content Warning and Disclaimer:<\/strong> This chapter includes images of human remains as well as discussions centered on human skeletal analyses. All images are derived from casts, sketches, nonhuman skeletal material, as well as non-Indigenous skeletal materials curated within the CSU, Chico Human Identification Lab, and the Hartnett-Fulginiti donated skeletal collection.<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\"><span style=\"color: #ffffff\">Learning Objectives<\/span><\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Define forensic anthropology as a subfield of biological anthropology.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Describe the seven steps carried out during skeletal analysis.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Outline the four major components of the biological profile.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Contrast the four categories of trauma.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Explain how to identify the different taphonomic agents that alter bone.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Discuss ethical considerations for forensic anthropology.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1214\">Forensic anthropology<\/a><\/strong> is a subfield of biological anthropology and an applied area of anthropology. Forensic anthropologists use skeletal analysis to gain information about humans in the present or recent past, then they apply this information within a medicolegal context. This means that forensic anthropologists specifically conduct their analysis on recently deceased individuals (typically within the last 50 years) as part of investigations by law enforcement. Forensic anthropologists can assist law enforcement agencies in several different ways, including aiding in the identification of human remains whether they are complete, fragmentary, burned, scattered, or decomposed. Additionally, forensic anthropologists can help determine what happened to the deceased at or around the time of death as well as what processes acted on the body after death (e.g., whether the remains were scattered by animals, whether they were buried in the ground, or whether they remained on the surface as the soft tissue decomposed).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Many times, because of their expertise in identifying human skeletal remains, forensic anthropologists are called to help with outdoor search-and-recovery efforts, such as locating remains scattered across the surface or carefully excavating and documenting buried remains. In other cases, forensic anthropologists recover remains after natural disasters or accidents, such as fire scenes, and can help identify whether each bone belongs to a human or an animal. Forensic anthropology spans a wide scope of contexts involving the law, including incidences of mass disasters, genocide, and war crimes.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">A point that can be somewhat confusing for students is that although the term <em>forensic<\/em> is included in this subfield of biological anthropology, there are many forensic techniques that are not included in the subfield. Almost exclusively, forensic anthropology deals with skeletal analysis. While this can include the comparison of antemortem (before death) and postmortem (after death) radiographs to identify whether remains belong to a specific person, or using photographic superimposition of the cranium, it does not include analyses beyond the skeleton. For example, blood-spatter analysis, DNA analysis, fingerprints, and material evidence collection do not fall under the scope of forensic anthropology.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">So, what can forensic anthropologists glean from bones alone? Forensic anthropologists can address a number of questions about a human individual based on their skeletal remains. Some of those questions are as follows: How old was the person? Was the person biologically male or female? How tall was the person? What happened to the person at or around their time of death? Were they sick? The information from the skeletal analysis can then be matched with missing persons records, medical records, or dental records, aiding law enforcement agencies with identifications and investigations.<\/p>\n<h2 class=\"import-Normal\">Skeletal Analysis<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropology relies on skeletal analysis to reveal information about the deceased. <span style=\"background-color: #00ffff\">The methodology and approaches outlined below are specific to the United States.<\/span> Forensic anthropological methods differ depending on the country conducting an investigation. In the United States, there are typically seven steps or questions to the process:<\/p>\n<ul>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Is it bone?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Is it human?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Is it modern or archeological?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">How many individuals are present or what is the minimum number of individuals (MNI)?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Who is it?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Is there evidence of trauma before or around the time of death?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">What happened to the remains after death?<\/li>\n<\/ul>\n<h3 class=\"import-Normal\"><strong>Is It Bone?<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">One of the most important steps in any skeletal analysis starts with determining whether or not material suspected to be bone is in fact bone. Though it goes without saying that a forensic anthropologist would only carry out analysis on bone, this step is not always straightforward. Whole bones are relatively easy to identify, but determining whether or not something is bone becomes more challenging once it becomes fragmentary. As an example, in high heat such as that seen on fire scenes, bone can break into pieces. During a house fire with fatalities, firefighters watered down the burning home. After the fire was extinguished, the sheetrock (used to construct the walls of the home) was drenched and crumbled. The crumbled sheetrock was similar in color and form to burned, fragmented bone, therefore mistakable for human remains (Figure 15.1). Forensic anthropologists on scene were able to separate the bones from the construction material, helping to confirm the presence of bone and hence the presence of individual victims of the fire. In this case, forensic anthropologists were able to recognize the anatomical and layered structure of bone and were able to distinguish it from the uniform and unlayered structure of sheetrock.<\/p>\n<p class=\"import-Normal\"><strong><img class=\"aligncenter\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/06\/image3.png\" alt=\"Long rectangular sheetrock with exposed porous surface.\" width=\"182\" height=\"208\" \/><\/strong><\/p>\n<figure style=\"width: 372px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image6-1.png\" alt=\"Two examples of sheetrock with dried or burnt surfaces.\" width=\"372\" height=\"210\" \/><figcaption class=\"wp-caption-text\">Figure 15.1: Burned sheetrock used as building material appears similar to human bone but can be differentiated by the fact that it is the same density throughout. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Example of burned sheetrock (Figure 15.1)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">As demonstrated by the example above, both the macrostructure (visible with the naked eye) and microstructure (visible with a microscope) of bone are helpful in bone identification. Bones are organs in the body made up of connective tissue. The connective tissue is hardened by a mineral deposition, which is why bone is rigid in comparison to other connective tissues such as cartilage (Tersigni-Tarrant and Langley 2017, 82\u201383; White and Folkens 2005, 31). In a living body, the mineralized tissue does not make up the only component of bone\u2014there are also blood, bone marrow, cartilage, and other types of tissues. However, in dry bone, two distinct layers of the bone are the most helpful for identification. The outer layer is made up of densely arranged osseous (bone) tissue called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1216\">compact (cortical) bone<\/a><\/strong>. The inner layer is composed of much more loosely organized, porous bone tissue whose appearance resembles that of a sponge, hence the name <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1218\"><strong>spongy (trabecular) bone<\/strong><\/a>. Knowing that most bone contains both layers helps with the macroscopic identification of bone (Figures 15.2, 15.3). For example, a piece of coconut shell might look a lot like a fragment of a human skull bone. However, closer inspection will demonstrate that coconut shell only has one very dense layer, while bone has both the compact and spongy layers.<\/p>\n<figure style=\"width: 380px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image27-1.png\" alt=\"Drawing showing thick exterior compact bone and porous internal cortical bone.\" width=\"380\" height=\"371\" \/><figcaption class=\"wp-caption-text\">Figure 15.2: Cross section of human long bone with compact and cortical bone layers visible. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Cross section of human long bone (Figure 15.2)<\/a> original to<a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"> Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p>&nbsp;<\/p>\n<figure style=\"width: 364px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image25-2.png\" alt=\"Cranial bone cross section called a periosteum with spongy bone (diploe) and compact bone labeled. Compact bone is a thin slice at the top and bottom and is smooth and hard. Spongy bone is in the middle and has irregular holes and indentations throughout. \" width=\"364\" height=\"184\" \/><figcaption class=\"wp-caption-text\">Figure 15.3: Cranial anatomy is slightly different as compared to that of a long bone in cross section. The compact (cortical) bone layers sandwich the spongy (trabecular) bone. One layer of compact bone forms the very outer surface of the skull and the other lines the internal surface of the skull. Credit: <a href=\"https:\/\/cnx.org\/contents\/FPtK1zmh@6.27:kwbeYj9S@3\/Bone-Structure\">Anatomy of a Flat Bone (Anatomy &amp; Physiology, Figure 6.3.3)<\/a> by<a href=\"https:\/\/openstax.org\/\"> OpenStax<\/a> is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/\"> CC BY 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The microscopic identification of bone relies on knowledge of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1220\">osteons<\/a><\/strong>, or bone cells (Figure 15.4). Under magnification, bone cells are visible in the outer, compact layer of bone. The bone cells are arranged in a concentric pattern around blood vessels for blood supply. The specific shape of the cells can help differentiate, for example, a small piece of PVC (white plastic) pipe from a human bone fragment (Figure 15.5).<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 340px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image14-3.png\" alt=\"Microscope image showing clustered osteons. Each has many rings and a dark center.\" width=\"340\" height=\"218\" \/><figcaption class=\"wp-caption-text\">Figure 15.4: Bone microstructure (osteons). Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Bone_(248_12)_Bone_cross_section.jpg\">Bone (248 12) Bone cross section<\/a> by <a href=\"https:\/\/cs.wikipedia.org\/wiki\/Josef_Reischig\">Doc. RNDr. Josef Reischig, CSc.<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p>&nbsp;<\/p>\n<figure style=\"width: 332px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image7-1.png\" alt=\"Flat, white section of PVC. Edges are broken and surface rough.\" width=\"332\" height=\"268\" \/><figcaption class=\"wp-caption-text\">Figure 15.5: Fragments of plastic PVC pipe, such as those seen in this photo, may be mistaken for human bone. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Example of PVC pipe<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Is It Human?<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Once it has been determined that an object is bone, the next logical step is to identify whether the bone belongs to a human or an animal. Forensic anthropologists are faced with this question in everyday practice because human versus nonhuman bone identification is one of the most frequent requests they receive from law enforcement agencies.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">There are many different ways to distinguish human versus nonhuman bone. The morphology (the shape\/form) of human bone is a good place for students to start. Identifying the 206 bones in the adult human skeleton and each bone\u2019s distinguishing features (muscle attachment sites, openings and grooves for nerves and blood vessels, etc.) is fundamental to skeletal analysis.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Nevertheless, there are many animal bones and human bones that look similar. For example, the declawed skeleton of a bear paw looks a lot like a human hand, pig molars appear similar to human molars, and some smaller animal bones might be mistaken for those of an infant. To add to the confusion, fragmentary bone may be even more difficult to identify as human or nonhuman. However, several major differences between human and nonhuman vertebrate bone help distinguish the two.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropologists pay special attention to the density of the outer, compact layer of bone in both the cranium and in the long bones. Human cranial bone has three distinctive layers. The spongy bone is sandwiched between the outer (ectocranial) and inner (endocranial) compact layers. In most other mammals, the distinction between the spongy and compact layers is not always so definite. Secondly, the compact layer in nonhuman mammal long bones can be much thicker than observed in human bone. Due to the increased density of the compact layer, nonhuman bone tends to be heavier than human bone (Figure 15.6).<\/p>\n<figure style=\"width: 399px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image11-2.png\" alt=\"Ring-like cross section of bone.\" width=\"399\" height=\"266\" \/><figcaption class=\"wp-caption-text\">Figure 15.6: The compact layer of this animal bone is very thick, with almost no spongy bone visible. Compare with Figure 15.2 to visualize the difference in structure between human and nonhuman bone. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Animal bone cross section (Figure 15.6)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The size of a bone can also help determine whether it belongs to a human. Adult human bones are larger than subadult or infant bones. However, another major difference between human adult bones and those of a young individual or infant human can be attributed to development and growth of the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1222\">epiphyses<\/a><\/strong> (ends of the bone). The epiphyses of human subadult bones are not fused to the shaft (Figure 15.7). Therefore, if a bone is small and it is suspected to belong to a human subadult or infant, the epiphyses would not be fused. Many small animal bones appear very similar in form compared to adult human bones, but they are much too small to belong to an adult human. Yet they can be eliminated as subadult or infant bones if the epiphyses are fused to the shaft.<\/p>\n<figure style=\"width: 288px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image13-3.png\" alt=\"X-ray image of child\u2019s ankle.\" width=\"288\" height=\"412\" \/><figcaption class=\"wp-caption-text\">Figure 15.7: An x-ray of a subadult\u2019s ankle with the epiphyses of the tibia and fibula visible. The gap between the shaft of the bone and the end of the bone (epiphysis) is the location of the growth plate. Therefore, the growth plate gap is what separates the shafts from the epiphyses in the image. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Tib_fib_growth_plates.jpg\">Tib fib growth plates<\/a> by <a href=\"https:\/\/en.wikipedia.org\/wiki\/User:Gilo1969\">Gilo1969<\/a> at <a href=\"https:\/\/en.wikipedia.org\/wiki\/\">English Wikipedia<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/3.0\/legalcode\">CC BY 3.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Is It Modern or Archaeological? <\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropologists work with modern cases that fall within the scope of law enforcement investigations. Accordingly, it is important to determine whether discovered human remains are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1224\">archaeological<\/a> <\/strong>or forensic in nature. Human remains that are historic are considered archeaological. The scientific study of human remains from archaeological sites is called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1226\">bioarchaeology<\/a><\/strong>.<\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Dig Deeper: Bioarchaeology<\/h2>\n<p class=\"import-Normal\">For readers who are interested in the sister subfield of bioarchaeology, which studies human remains and material culture from the past, please refer to chapter 8 of <em>Bioarchaeology: Interpreting Human Behavior from Skeletal Remains,<\/em> in <em>TRACES: An Open Invitation to Archaeology<\/em> (Blatt, Michael, and Bright forthcoming).<\/p>\n<\/div>\n<p>A forensic anthropologist should begin their analysis by reviewing the context in which the remains were discovered. This will help them understand a great deal about the remains, including determining whether they are archaeological or forensic in nature as well as considering legal and ethical issues associated with the collection, analysis, and storage of human remains (see \u201cEthics and Human Rights\u201d section of this chapter for more information).<\/p>\n<figure style=\"width: 403px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image10-3.png\" alt=\"Four teeth in a person\u2019s mouth. First molar with silver filling.\" width=\"403\" height=\"303\" \/><figcaption class=\"wp-caption-text\">Figure 15.8: A human tooth with a filling. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Filling.jpg#filehistory\">Filling<\/a> by Kauzio has been designated to the <a href=\"https:\/\/creativecommons.org\/share-your-work\/public-domain\/cc0\/\">public domain (CC0)<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The \u201ccontext\u201d refers to the relationship the remains have to the immediate area in which they were found. This includes the specific place where the remains were found, the soil or other organic matter immediately surrounding the remains, and any other objects or artifacts in close proximity to the body. For example, imagine that a set of remains has been located during a house renovation. The remains are discovered below the foundation. Do the remains belong to a murder victim? Or was the house built on top of an ancient burial ground? Observing information from the surroundings can help determine whether the remains are archaeological or modern. How long ago was the foundation of the house erected? Are there artifacts in close proximity to the body, such as clothing or stone tools? These are questions about the surroundings that will help determine the relative age of the remains.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Clues directly from the skeleton may also indicate whether the remains are archaeological or modern. For example, tooth fillings can suggest that the individual was alive recently (Figure 15.8). In fact, filling material has changed over the decades, so the specific type of material used to fix a cavity can be matched with specific time periods. Gold was used in dental work in the past, but more recently composite (a mixture of plastic and fine glass) fillings have become more common.<\/p>\n<h3><strong>How <\/strong><strong>Many Individuals Are Present?<\/strong><\/h3>\n<h4 class=\"import-Normal\"><em>What Is MNI?<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Another assessment that an anthropologist can perform is the calculation of the number of individuals in a mixed burial assemblage. Because not all burials consist of a single individual, it is important to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1268\">burial assemblage<\/a><\/strong> be able to estimate the number of individuals in a forensic context. Quantification of the number of individuals in a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1524\">burial assemblage<\/a><\/strong> can be done through the application of a number of methods, including the following: the Minimum Number of Individuals (MNI), the Most Likely Number of Individuals (MLNI), and the Lincoln Index (LI). The most commonly used method in biological anthropology, and the focus of this section, is determination of the MNI.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The MNI presents \u201cthe minimum estimate for the number of individuals that contributed to the sample\u201d (Adams and Konigsberg 2008, 243). Many methods of calculating MNI were originally developed within the field of zooarchaeology for use on calculating the number of individuals in faunal or animal assemblages (Adams and Konigsberg 2008, 241). What MNI calculations provide is a lowest possible count for the total number of individuals contributing to a skeletal assemblage. Traditional methods of calculating MNI include separating a skeletal assemblage into categories according to the individual bone and the side the bone comes from and then taking the highest count per category and assigning that as the minimum number (Figure 15.9).<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 664px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image28-3.png\" alt=\"Many bone portions laying on individual plastic bags on a table.\" width=\"664\" height=\"441\" \/><figcaption class=\"wp-caption-text\">Figure 15.9: Skeletal elements from a commingled faunal assemblage. Credit: Commingled animal remains from Eden-Farson Pre-Contact site in southwest Wyoming by Matt O\u2019Brien original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Why Calculate MNI?<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In a forensic context, the determination of MNI is most applicable in cases of mass graves, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1232\">commingled burials<\/a><\/strong>, and mass fatality incidents. The term <em>commingled<\/em> is applied to any burial assemblage in which individual skeletons are not separated into separate burials. As an example, the authors of this chapter have observed commingling of remains resulting from mass fatality wildfire events. Commingled remains may also be encountered in events such as a plane or vehicle crash. It is important to remember that in any forensic context, MNI should be referenced and an MNI of one should be substantiated by the fact that there was no repetition of elements associated with the case.<\/p>\n<h3 class=\"import-Normal\"><strong>Constructing the Biological Profile<\/strong><\/h3>\n<h4 class=\"import-Normal\"><em>Who Is It?<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">\u201cWho is it?\u201d is one of the first questions that law enforcement officers ask when they are faced with a set of skeletal remains. To answer this question, forensic anthropologists construct a biological profile (White and Folkens 2005, 405). A <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1228\">biological profile<\/a> <\/strong>is an individual\u2019s identifying characteristics, or biological information, which include the following: biological sex, age at death, stature, population affinity, skeletal variation, and evidence of trauma and pathology.<\/p>\n<h4 class=\"import-Normal\"><em>Assessing Biological Sex <\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Assessment of biological sex is often one of the first things considered when establishing a biological profile because several other parts, such as age and stature estimations, rely on an assessment of biological sex to make the calculations more accurate.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Assessment of biological sex focuses on differences in both morphological (form or structure) and metric (measured) traits in individuals. When assessing morphological traits, the skull and the pelvis are the most commonly referenced areas of the skeleton. These differences are related to sexual dimorphism usually varying in the amount of robusticity seen between males and females. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1230\">Robusticity<\/a> <\/strong>deals with strength and size; it is frequently used as a term to describe a large size or thickness. In general, males will show a greater degree of robusticity than females. For example, the length and width of the mastoid process, a bony projection located behind the opening for the ear, is typically larger in males. The mastoid process is an attachment point for muscles of the neck, and this bony projection tends to be wider and longer in males. In general, cranial features tend to be more robust in males (Figure 15.10).<\/p>\n<figure style=\"width: 601px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image26-3.png\" alt=\"Front and side images of a male (left) and female (right) cranium.\" width=\"601\" height=\"632\" \/><figcaption class=\"wp-caption-text\">Figure 15.10: Anterior and lateral view of a male and female cranium. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Anterior and lateral view of a male and female cranium (Figure 15.10)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropo logy<\/a> by Ashley Kendell is a collective work under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Includes <a href=\"https:\/\/boneclones.com\/product\/modern-human-asian-female-skull-BC-149\/category\/all-human-skulls\/human-anatomy\">Human Female Asian Skull<\/a> and <a href=\"https:\/\/boneclones.com\/product\/human-asian-male-skull-BC-016\/category\/all-human-skulls\/human-anatomy\">Human Male Asian Skull<\/a> by <a href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a>, used by permission.]<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">When considering the pelvis, the features associated with the ability to give birth help distinguish females from males. During puberty, estrogen causes a widening of the female pelvis to allow for the passage of a baby. Several studies have identified specific features or bony landmarks associated with the widening of the hips, and this section will discuss one such method. The Phenice Method (Phenice 1969) is traditionally the most common reference used to assess morphological characteristics associated with sex. The Phenice Method specifically looks at the presence or absence of (1) a ventral arc, (2) the presence or absence of a subpubic concavity, and (3) the width of the medial aspect of the ischiopubic ramus (Figure 15.11). When present, the ventral arc, a ridge of bone located on the ventral surface of the pubic bone, is indicative of female remains. Likewise the presence of a subpubic concavity and a narrow medial aspect of the ischiopubic ramus is associated with a female sex estimation. Assessments of these features, as well as those of the skull (when both the pelvis and skull are present), are combined for an overall estimation of sex.<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 1603px\" class=\"wp-caption alignnone\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image29-3.png\" alt=\"Male and female os coxae (anterior portions).\" width=\"1603\" height=\"582\" \/><figcaption class=\"wp-caption-text\">Figure 15.11: Features associated with the Phenice Method. Images derived from CSU-HIL donated skeletal collection. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Features associated with the Phenice Method (Figure 15.11)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Colleen Milligan is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Metric analyses are also used in the estimation of sex. Measurements taken from every region of the body can contribute to estimating sex through statistical approaches that assign a predictive value of sex. These approaches can include multiple measurements from several skeletal elements in what is called multivariate (multiple variables) statistics. Other approaches consider a single measurement, such as the diameter of the head of the femur, of a specific element in a univariate (single variable) analysis (Berg 2017, 152\u2013156).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">It is important to note that, although forensic anthropologists usually begin assessment of biological profile with biological sex, there is one major instance in which this is not appropriate. The case of two individuals found in California, on July 8, 1979, is one example that demonstrates the effect age has on the estimation of sex. The identities of the two individuals were unknown; therefore, law enforcement sent them to a lab for identification. A skeletal analysis determined that the remains represented one adolescent male and one adolescent female, both younger than 18 years of age. This information did not match with any known missing children at the time.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In 2015, the cold case was reanalyzed, and DNA samples were extracted. The results indicated that the remains were actually those of two girls who went missing in 1978. The girls were 15 years old and 14 years old at the time of death. It is clear that the 1979 results were incorrect, but this mistake also provides the opportunity to discuss the limitations of assessing sex from a subadult skeleton.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Assessing sex from the human skeleton is based on biological and genetic traits associated with females and males. These traits are linked to differences in sexual dimorphism and reproductive characteristics between females and males. The link to reproductive characteristics means that most indicators of biological sex do not fully manifest in prepubescent individuals, making estimations of sex unreliable in younger individuals (SWGANTH 2010b). This was the case in the example of the 14-year-old girl. When examined in 1979, her remains were misidentified as male because she had not yet fully developed female pelvic traits.<\/p>\n<h4 class=\"import-Normal\"><em>Sex vs. Gender<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Biological sex is a different concept than <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1234\">gender<\/a><\/strong>. While biological anthropologists can estimate sex from the skeleton, estimating an individual\u2019s gender would require a greater context because gender is defined culturally rather than biologically. Take, for example, an individual who identifies as transgender. This individual has a gender identity that is different from their biological sex. The gender identity of any individual depends on factors related to self-identification, situation or context, and cultural factors. <span style=\"background-color: #00ffff\">While in the U.S<\/span>. we have historically thought of sex and gender as binary concepts (male or female), many cultures throughout the world recognize several possible gender identities. In this sense, gender is seen as a continuous or fluid variable rather than a fixed one.<\/p>\n<p class=\"import-Normal\">Historically, forensic anthropologists have used a binary construct to categorize human skeletal remains as either male or female (with the accompanying categories of probable male, probable female, and indeterminate). In the case of transgender and gender nonconforming individuals, the binary approach to sex assessment may delay or hinder identification efforts (Buchanan 2014; Schall, Rogers, and Deschamps-Braly 2020; Tallman, Kincer, and Plemons 2021). As such, many forensic anthropologists have begun to address the inherent problems associated with a binary approach to sex identification and to explore ways of assessing social identity and self-identified gender using skeletal remains and forensic context.<\/p>\n<p class=\"import-Normal\">For the duration of this section, the term <em>transgender<\/em> refers to individuals whose gender identity differs from the sex assigned at birth (Schall, Rogers, and Deschamps-Braly 2020:2). Transgender individuals transition from one gender binary to another, such as male-to-female (MTF) or female-to-male (FTM). While many of the gender-affirming procedures available to trans and gender-nonconforming individuals are focused on soft tissue modifications (e.g., breast augmentation, genital reconstruction, hormone therapies, etc.), there are a number of gender-affirmation surgeries that do leave a permanent record on the skeleton. Generally speaking, FTM transgender people are reported to undergo fewer surgical procedures than do MTF transgender people (Buchanan 2014). The discussion below focuses on Facial Feminization Surgery (FFS), which leaves a permanent record on the human skeleton that may be used to help make an identification.<\/p>\n<p class=\"import-Normal\">FFS refers to a combination of procedures focused on sexually dimorphic features of the face, with the intent of transforming typically male facial features into more feminine forms. Facial Feminization Surgery procedures were developed by Dr. Douglas Ousterhout, a San Francisco based cranio-maxillofacial surgeon, in the mid-1980s (Schall, Rogers, and Deschamps-Braly 2020:2). FFS can include one or a combination of the following: hairline lowering, forehead reduction and contouring, brow lift, reduction rhinoplasty, cheek enhancement, lift lift, lip filling, chin contouring, jaw contouring, and\/or tracheal shave (Buchanan 2014; Schall, Rogers, and Deschamps-Braly 2020:2). Of the procedures outlined previously, four are known to directly affect the facial skeleton: forehead contouring, rhinoplasty, chin contouring, and jaw contouring (Buchanan 2014; Schall, Rogers, and Deschamps-Braly 2020:2).<\/p>\n<p class=\"import-Normal\">Because FFS procedures have been widely documented in the medical (and more recently the forensic anthropological) literature, there are a number of indicators that a forensic anthropologist can use to make more informed evaluations of gender, including evidence of bone remodeling in sexually dimorphic regions of the skull (e.g., forehead, chin, jawline), as well as the presence of plates, pins, or other surgical hardware that may be evidence of FFS (Buchanan 2014; Schall, Rogers, and Deschamps-Braly 2020; Tallman, Kincer, and Plemons 2021). Additionally, some forensic anthropologists suggest cautiously integrating contextual information from the scene, such as personal effects, material evidence, and recovery scene information, into their evaluation of an individual\u2019s social identity (Beatrice and Soler 2016; Birkby, Fenton, and Anderson 2008; Soler and Beatrice 2018; Soler et al. 2019; Tallman, Kincer, and Plemons 2021; Winburn, Schoff, and Warren 2016). The ultimate goal of many skeletal analyses is to make a positive identification on a set of unidentified remains.<\/p>\n<h4 class=\"import-Normal\"><em>Assessment <\/em><em>of Population Affinity<\/em><\/h4>\n<p>In an effort to combat the erroneous assumptions tied to the race concept, forensic anthropologists have attempted to reframe this component of the biological profile. The term <em>race<\/em> is no longer used in casework and teaching. Historically, the word <em>ancestry<\/em> is and was deemed a more appropriate way to describe an individual\u2019s phenotype. However, in more recent years, forensic anthropologists have begun using the term <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1236\">population affinity<\/a><\/strong><em>, <\/em>recognizing that we are basing our analysis on the similarities we see based on the reference samples we have available (Winburn and Algee-Hewitt 2021). An important note here is that it is possible to hinder identifications and harm individuals when tools like estimations of population affinity are misapplied, misinterpreted, or misused. For this reason, the field of forensic anthropology has ongoing conversations about the appropriateness of this analysis in the biological profile (Bethard and DiGangi 2020; Stull et al. 2021).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">We use the term <em>population affinity<\/em> to refer to the variation seen among modern populations\u2014variation that is both genetic and environmentally driven. The word <em>affinity<\/em> refers to similarities or relationships between individuals. As forensic anthropologists, we compare an unknown individual to multiple reference groups and look for the degree of similarity in observable traits with those groups. As noted previously, population affinity can aid law enforcement in their identification of missing persons or unknown skeletal remains.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Within the field of anthropology, the estimation of population affinity has a contentious history, and early attempts at classification were largely based on the erroneous assumption that an individual\u2019s <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1238\">phenotype <\/a><\/strong> (outward appearance) was correlated with their innate intelligence and abilities (see Chapter 13 for a more in-depth discussion of the history of the race concept). The use of the term <em>race<\/em> is deeply embedded in the social context of the United States. In any other organism\/living thing, groups divided according to the biological race concept would be defined as a separate subspecies. The major issue with applying the biological race concept to humans is that there are not enough differences between any two populations to separate on a genetic basis. In other words, <em>biological races do not exist in human populations. <\/em>However, the concept of race has been perpetuated and upheld by sociocultural constructs of race.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The conundrum for forensic anthropologists is the fact that while races do not exist on a biological level, we still socially recognize and categorize individuals based on their phenotype. Clearly, our phenotype is an important factor in not only how we are viewed by others but also how we identify ourselves. It is also a commonly reported variable. Often labeled as \u201crace,\u201d we are asked to report how we self-identify on school applications, government identification, surveys, census reports, and so forth. It follows then that when a person is reported missing, the information commonly collected by law enforcement and sometimes entered into a missing person\u2019s database includes their age, biological sex, stature, and \u201crace.\u201d Therefore, the more information a forensic anthropologist can provide regarding the individual\u2019s physical characteristics, the more he or she can help to narrow the search.<\/p>\n<p class=\"import-Normal\">As an exercise, create a list of all of the women you know who are between the ages of 18 and 24 and approximately 5\u2019 4\u201d to 5\u2019 9\u201d tall. You probably have several dozen people on the list. Now, consider how many females you know who are between the ages of 18 and 24, are approximately 5\u2019 4\u201d to 5\u2019 9\u201d tall, and are Vietnamese. Your list is going to be significantly shorter. That\u2019s how missing persons searches go as well. The more information you can provide regarding a decedent\u2019s phenotype, the fewer possible matches law enforcement are left to investigate. This is why population affinity has historically been included as a part of the biological profile.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Traditionally, population affinity was accomplished through a visual inspection of morphological variants of the skull (morphoscopics). These methods focused on elements of the facial skeleton, including the nose, eyes, and cheek bones. However, in an effort to reduce subjectivity, nonmetric cranial traits are now assessed within a statistical framework to help anthropologists better interpret their distribution among living populations (Hefner and Linde 2018). Based on the observable traits, a macromorphoscopic analysis will allow the practitioner to create a statistical prediction of geographic origin. In essence, forensic anthropologists are using human variation in the estimation of geographic origin, by referencing documented frequencies of nonmetric skeletal indicators or macromorphoscopic traits.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Population affinity is also assessed through metric analyses. The computer program Fordisc is an anthropological tool used to estimate different components of the biological profile, including ancestry, sex, and stature. When using Fordisc, skeletal measurements are input into the computer software, and the program employs multivariate statistical classification methods, including discriminant function analysis, to generate a statistical prediction for the geographic origin of unknown remains based on the comparison of the unknown to the reference samples in the software program. Fordisc also calculates the likelihood of the prediction being correct, as well as how typical the metric data is for the assigned group.<\/p>\n<h4 class=\"import-Normal\"><em>Estimating Age-at-Death<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Estimating age-at-death from the skeleton relies on the measurement of two basic physiological processes: (1) growth and development and (2) degeneration (or aging). From fetal development on, our bones and teeth grow and change at a predictable rate. This provides for relatively accurate age estimates. After our bones and teeth cease to grow and develop, they begin to undergo structural changes, or degeneration, associated with aging. This does not happen at such predictable rates and, therefore, results in less accurate or larger age-range estimations.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">During growth and development stages, two primary methods used for estimations of age of subadults (those under the age of 18) are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1240\">epiphyseal union<\/a><\/strong> and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1244\">dental development.<\/a><\/strong> Epiphyseal union<strong> (<\/strong>or <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1242\">epiphyseal fusion<\/a><\/strong>) refers to the appearance and closure of the epiphyseal plates between the primary centers of growth in a bone and the subsequent centers of growth (see Figure 15.7). Prior to complete union, the cartilaginous area between the primary and secondary centers of growth is also referred to as the growth plates (Schaefer, Black, and Scheuer 2009). Different areas of the skeleton have documented differences in the appearance and closure of epiphyses, making this a reliable method for aging subadult remains (SWGANTH 2013).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">As an example of its utility in the identification process, epiphyseal development was used to identify two subadult victims of a fatal fire in Flint, Michigan, in February 2010. The remains represented two young girls, ages three and four. Due to the intensity of the fire, the subadult victims were differentiated from each other through the appearance of the patella, the kneecap. The patella is a bone that develops within the tendon of the quadriceps muscle at the knee joint. The patella begins to form around three to four years of age (Cunningham, Scheuer, and Black 2016, 407\u2013409). In the example above, radiographs of the knees showed the presence of a patella in the four-year-old girl and the absence of a clearly discernible patella in the three-year-old.<\/p>\n<figure style=\"width: 358px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image18-2.png\" alt=\"Cranial cast of child with exposed maxilla and mandible to see developing dentition.\" width=\"358\" height=\"358\" \/><figcaption class=\"wp-caption-text\">Figure 15.12: Dental development in a subadult. Credit: <a href=\"https:\/\/boneclones.com\/product\/5-year-old-human-child-skull-with-mixed-dentition-exposed-BC-189\">5-year-old Human Child Skull with Mixed Dentition Exposed<\/a> by <a href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a> is used by permission and available here is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Dental development begins during fetal stages of growth and continues until the complete formation and eruption of the adult third molars (if present). The first set of teeth to appear are called deciduous or baby teeth. Individuals develop a total of 20 deciduous teeth, including incisors, canines, and molars. These are generally replaced by adult dentition as an individual grows (Figure 15.12). A total of 32 teeth are represented in the adult dental arcade, including incisors, canines, premolars, and molars. When dental development is used for age estimations, researchers use both tooth-formation patterns and eruption schedules as determining evidence. For example, the crown of the tooth forms first followed by the formation of the tooth root. During development, an individual can exhibit a partially formed crown or a complete crown with a partially formed root. The teeth generally begin the eruption process once the crown of the tooth is complete. The developmental stages of dentition are one of the most reliable and consistent aging methods for subadults (Langley, Gooding, and Tersigni-Tarrant 2017, 176\u2013177).<\/p>\n<figure style=\"width: 403px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image22-3.png\" alt=\"Surfaces of three pubic symphyses: billowy (A) to more flat (B) to rough (C).\" width=\"403\" height=\"224\" \/><figcaption class=\"wp-caption-text\">Figure 15.13: Examples of degenerative changes to the pubic symphysis: (A) young adult; (B) middle adult; (C) old adult. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Example of the progression of degenerative changes to the pubic symphysis (Figure 15.14)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropo logy<\/a> by Ashley Kendell is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Original photos by Dr. Julie Fleischman used by permission. Pubic symphyses are curated in the Hartnett-Fulginiti donated skeletal collection. Donation and research consent was provided by next of kin.]<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Degenerative changes in the skeleton typically begin after 18 years of age, with more prominent changes developing after an individual reaches middle adulthood (commonly defined as after 35 years of age in osteology). These changes are most easily seen around joint surfaces of the pelvis, the cranial vault, and the ribs. In this chapter, we focus on the pubic symphysis surfaces of the pelvis and the sternal ends of the ribs, which show metamorphic changes from young adulthood to older adulthood. The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1262\">pubic symphysis<\/a> <\/strong>is a joint that unites the left and right halves of the pelvis. The surface of the pubic symphysis changes during adulthood, beginning as a surface with pronounced ridges (called billowing) and flattening with a more distinct rim to the pubic symphysis as an individual ages. As with all metamorphic age changes, older adults tend to develop lipping around the joint surfaces as well as a breakdown of the joint surfaces. The most commonly used method for aging adult skeletons from the pubic symphysis is the Suchey-Brooks method (Brooks and Suchey 1990; Katz and Suchey 1986). This method divides the changes seen with the pubic symphysis into six phases based on macroscopic age-related changes to the surface. Figure 15.13 provides a visual of the degenerative changes that typically occur on the pubic symphysis.<\/p>\n<figure style=\"width: 403px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image12-3.png\" alt=\"Three sternal rib ends demonstrating progressive changes that occur with age.\" width=\"403\" height=\"220\" \/><figcaption class=\"wp-caption-text\">Figure 15.14: Examples of degenerative changes to the sternal rib end: (A) young adult; (B) middle adult; (C) old adult. Images derived from CSU, Chico HIL donated skeletal collection. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Examples of degenerative changes to the sternal rib end (Figure 15.15)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The sternal end of the ribs, the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1246\">anterior <\/a><\/strong> end of the rib that connects via cartilage to the sternum, is also used in age estimations of adults. This method, first developed by M. Y. \u0130\u015fcan and colleagues, considers both the change in shape of the sternal end as well as the quality of the bone (\u0130\u015fcan, Loth, and Wright 1984; \u0130\u015fcan, Loth, and Wright 1985). The sternal end first develops a billowing appearance in young adulthood. The bone typically develops a wider and deeper cupped end as an individual ages. Older adults tend to exhibit bony extensions of the sternal end rim as attaching cartilage ossifies. Figure 15.14 provides a visual of the degenerative changes that typically occur in sternal rib ends.<\/p>\n<h4 class=\"import-Normal\"><em>Estimating Stature<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Stature, or height, is one of the most prominently recorded components of the biological profile. Our height is recorded from infancy through adulthood. Doctor\u2019s appointments, driver's license applications, and sports rosters all typically involve a measure of stature for an individual. As such, it is also a component of the biological profile nearly every individual will have on record. Bioarchaeologists and forensic anthropologists use stature estimation methods to provide a range within which an individual\u2019s biological height would fall. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1248\">Biological height<\/a> <\/strong>is a person\u2019s true anatomical height. However, the range created through these estimations is often compared to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1264\">reported stature<\/a><\/strong>, which is typically self-reported and based on an approximation of an individual\u2019s true height (Ousley 1995).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In June 2015, two men were shot and killed in Granite Bay, California, in a double homicide. Investigators were able to locate surveillance camera footage from a gas station where the two victims were spotted in a car with another individual believed to be the perpetrator in the case. The suspect, sitting behind the victims in the car, hung his right arm out of the window as the car drove away. The search for the perpetrator was eventually narrowed down to two suspects. One suspect was 5\u2019 8\u201d while the other suspect was 6\u2019 4\u201d, representing almost a foot difference in height reported stature between the two. Forensic anthropologists were given the dimensions of the car (for proportionality of the arm) and were asked to calculate the stature of the suspect in the car from measurements of the suspect\u2019s forearm hanging from the window. Approximate lengths of the bones of the forearm were established from the video footage and used to create a predicted stature range. Stature estimations from skeletal remains typically look at the correlation between the measurements of any individual bone and the overall measurement of body height. In the case above, the length of the right forearm pointed to the taller of the two suspects who was subsequently arrested for the homicide.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Certain bones, such as the long bones of the leg, contribute more to our overall height than others and can be used with mathematical equations known as regression equations. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1252\">Regression methods <\/a> <\/strong>examine the relationship between variables such as height and bone length and use the correlation between the variables to create a prediction interval (or range) for estimated stature. This method for calculating stature is the most commonly used method (SWGANTH 2012). Figure 15.15 shows the measurement of the bicondylar length of the femur for stature estimations.<\/p>\n<figure style=\"width: 584px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image2-3.png\" alt=\"A femur is measured using a wooden osteometric board.\" width=\"584\" height=\"389\" \/><figcaption class=\"wp-caption-text\">Figure 15.15: Image of measurement of the bicondylar length of the femur, often used in the estimation of living stature. Image derived from CSU, Chico HIL donated skeletal collection. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Measurement of the bicondylar length of the femur<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Identification Using Individualizing Characteristics<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">One of the most frequently requested analyses within the forensic anthropology laboratory is assistance with the identification of unidentified remains. While all components of a biological profile, as discussed above, can assist law enforcement officers and medical examiners to narrow down the list of potential identifications, a biological profile will not lead to a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1254\">positive identification<\/a><\/strong>. The term <em>positive identification<\/em> refers to a scientifically validated method of identifying previously unidentified remains. Presumptive identifications, however, are not scientifically validated; rather, they are based on circumstances or scene context. For example, if a decedent is found in a locked home with no evidence of forced entry but the body is no longer visually identifiable, it may be presumed that the remains belong to the homeowner. Hence, a presumptive identification.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The medicolegal system ultimately requires that a positive identification be made in such circumstances, and a presumptive identification is often a good way to narrow down the pool of possibilities. Biological profile information also assists with making a presumptive identification based on an individual\u2019s phenotype in life (e.g., what they looked like). As an example, a forensic anthropologist may establish the following components of a biological profile: white male, between the ages of 35 and 50, approximately 5\u2019 7\u201d to 5\u2019 11.\u201d While this seems like a rather specific description of an individual, you can imagine that this description fits dozens, if not hundreds, of people in an urban area. Therefore, law enforcement can use the biological profile information to narrow their pool of possible identifications to include only white males who fit the age and height outlined above. Once a possible match is found, the decedent can be identified using a method of positive identification.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Positive identifications are based on what we refer to as individualizing traits or characteristics, which are traits that are unique at the individual level. For example, brown hair is not an individualizing trait as brown is the most common hair color in the U.S. But, a specific pattern of dental restorations or surgical implants can be individualizing, because it is unlikely that you will have an exact match on either of these traits when comparing two individuals.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">A number of positive methods are available to forensic anthropologists, and for the remainder of this section we will discuss the following methods: comparative medical and dental radiography and identification of surgical implants.<\/p>\n<figure style=\"width: 165px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image17-3.png\" alt=\"Radiograph of skull with frontal sinuses visible.\" width=\"165\" height=\"182\" \/><figcaption class=\"wp-caption-text\">Figure 15.16: Example of the unique shape of the frontal sinus. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Frontal_bone_sinuses.jpg\">Frontal bone sinuses<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Alex_Khimich\">Alex Khimich<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Comparative medical and dental radiography is used to find consistency of traits when comparing antemortem records (medical and dental records taken during life) with images taken postmortem (after death). Comparative medical radiography focuses primarily on features associated with the skeletal system, including trabecular pattern (internal structure of bone that is honeycomb in appearance), bone shape or cortical density (compact outer layer of bone), and evidence of past trauma, skeletal pathology, or skeletal anomalies. Other individualizing traits include the shape of various bones or their features, such as the frontal sinuses (Figure 15.16).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Comparative dental radiography focuses on the number, shape, location, and orientation of dentition and dental restorations in antemortem and postmortem images. While there is not a minimum number of matching traits that need to be identified for an identification to be made, the antemortem and postmortem records should have enough skeletal or dental consistencies to conclude that the records did in fact come from the same individual (SWGANTH 2010a). Consideration should also be given to population-level frequencies of specific skeletal and dental traits. If a trait is particularly common within a given population, it may not be a good trait to utilize for positive identification.<\/p>\n<figure style=\"width: 354px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image16-3.png\" alt=\"A scapula and humerus with a metal shoulder replacement.\" width=\"354\" height=\"231\" \/><figcaption class=\"wp-caption-text\">Figure 15.17: Image of joint replacement in the right shoulder. Credit: <a href=\"https:\/\/naturalhistory.si.edu\/education\/teaching-resources\/written-bone\/skeleton-keys\/todays-bones\">Shoulder replacement<\/a> by <a href=\"https:\/\/www.si.edu\/\">Smithsonian<\/a> [exhibit: Written in Bone, Today\u2019s Bones] <a href=\"https:\/\/www.si.edu\/termsofuse\">is used for educational and non-commercial purposes as outlined by the Smithsonian.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Surgical implants or devices can also be used for identification purposes (Figure 15.17). These implements are sometimes recovered with human remains. One of the ways forensic anthropologists can use surgical implants to assist in decedent identification is by providing a thorough analysis of the implant and noting any identifying information such as serial numbers, manufacturer symbols, and so forth. This information can then sometimes be tracked directly to the manufacturer or the place of surgical intervention, which may be used to identify unknown remains (SWGANTH 2010a).<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Trans Doe Task Force<\/h2>\n<p class=\"import-Normal\">The Trans Doe Task Force (TDTF) is a Trans-led nonprofit organization that investigates cases involving LGBTQ+ missing and murdered persons. The organization specifically focuses on transgender and gender-variant cases, providing connections between law enforcement agencies, medical examiner offices, forensic anthropologists, and forensic genetic genealogists to increase the chances of identification. Additionally, the TDTF curates a data repository of missing, murdered, and unclaimed LGBTQ+ individuals, and they continuously try innovative approaches to identify these individuals, whose lived gender identity may not match their biological sex.<\/p>\n<p class=\"import-Normal\">For more information visit <a href=\"https:\/\/transdoetaskforce.org\/\">transdoetaskforce.org<\/a><\/p>\n<\/div>\n<h3 class=\"import-Normal\"><strong>Trauma Analysis<\/strong><\/h3>\n<h4 class=\"import-Normal\"><em>Types of Trauma<\/em><strong><br \/>\n<\/strong><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Within the field of anthropology, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1256\">trauma<\/a> <\/strong>is defined as an injury to living tissue caused by an extrinsic force or mechanism (Lovell 1997:139). Forensic anthropologists can assist a forensic pathologist by providing an interpretation of the course of events that led to skeletal trauma. Typically, traumatic injury to bone is classified into one of four categories, defined by the trauma mechanism. A trauma mechanism refers to the force that produced the skeletal modification and can be classified as (1) sharp force, (2) blunt force, (3) projectile, or (4) thermal (burning). Each type of trauma, and the characteristic pattern(s) associated with that particular categorization, will be discussed below.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">First, let\u2019s consider s<em>harp-force trauma<\/em>, which is caused by a tool that is edged, pointed, or beveled\u2014for example, a knife, saw, or machete (SWGANTH 2011). The patterns of injury resulting from sharp-force trauma include linear incisions created by a sharp, straight edge; punctures; and chop marks (Figure 15.18; SWGANTH 2011). When observed under a microscope, an anthropologist can often determine what kind of tool created the bone trauma. For example, a power saw cut will be discernible from a manual saw cut.<\/p>\n<figure style=\"width: 602px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image21-1.png\" alt=\"Anterior image of a skull with multiple traumatic injuries to forehead.\" width=\"602\" height=\"457\" \/><figcaption class=\"wp-caption-text\">Figure 15.18: Example of sharp-force trauma (sword wound) to the frontal bone. The skull appears sliced with thin lines in two places across the top of the skull. Credit: <a href=\"https:\/\/openverse.org\/image\/909d1b77-ad5f-4cda-be44-6d9b5fbf14b9\/\">Female skull injured by a medieval sword<\/a> by <a href=\"https:\/\/sketchfab.com\/provinciaal_depot_noordholland\">Provinciaal depot voor archeologie Noord-Holland<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY 4.0 License<\/a>. The original image is a 3D model that can be manipulated on the <a href=\"https:\/\/wordpress.org\/openverse\/image\/909d1b77-ad5f-4cda-be44-6d9b5fbf14b9\/\">openverse website<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Second, <em>blunt-force trauma<\/em> is defined as \u201ca relatively low-velocity impact over a relatively large surface area\u201d (Galloway 1999, 5). Blunt-force injuries can result from impacts from clubs, sticks, fists, and so forth. Blunt-force impacts typically leave an injury at the point of impact but can also lead to bending and deformation in other regions of the bone. Depressions, fractures, and deformation at and around the site of impact are all characteristics of blunt-force trauma (Figure 15.19). As with sharp-force trauma, an anthropologist attempts to interpret blunt-force injuries, providing information pertaining to the type of tool used, the direction of impact, the sequence of impacts, if more than one, and the amount of force applied.<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 578px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image30.png\" alt=\"Cranium with two blunt force impacts from a hammer.\" width=\"578\" height=\"803\" \/><figcaption class=\"wp-caption-text\">Figure 15.19: Example of multiple blunt force impacts to the left parietal and frontal bones. There is one hole in the skull with fractured bone around the edges. There are also multiple spots across the back of the skull with depressions of various sizes. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Skull_hammer_trauma.jpg\">Skull hammer trauma<\/a> by <a href=\"https:\/\/www.nih.gov\/\">the National Institutes of Health<\/a>, Health &amp; Human Services, is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>. [Exhibit: Visible Proofs: Forensic Views of the Body, U.S. National Library of Medicine, 19th Century Collection, National Museum of Health and Medicine, Armed Forces Institute of Pathology, Washington, D.C.]<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Third, <em>projectile trauma<\/em> refers to high-velocity trauma, typically affecting a small surface area (Galloway 1999, 6). Projectile trauma results from fast-moving objects such as bullets or shrapnel. It is typically characterized by penetrating defects or embedded materials (Figure 15.20). When interpreting injuries resulting from projectile trauma, an anthropologist can often offer information pertaining to the type of weapon used (e.g., rifle vs. handgun), relative size of the bullet (but not the caliber of the bullet), the direction the projectile was traveling, and the sequence of injuries if there are multiple present.<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 462px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image5-3.png\" alt=\"Anterior and posterior views of a skull with a gunshot wound.\" width=\"462\" height=\"291\" \/><figcaption class=\"wp-caption-text\">Figure 15.20: Example of projectile trauma with an entrance wound to the frontal bone and exit wound visible on the occipital. A small circular hole is visible in the front of the skull with cracks radiating out from the point of impact. There is a larger hole visible in the back of the skull that is irregular yet circular in shape. Credit: <a href=\"https:\/\/naturalhistory.si.edu\/education\/teaching-resources\/written-bone\/skeleton-keys\/how-bone-biographies-get-written\">Trauma: Gunshot Wounds<\/a> by <a href=\"https:\/\/www.si.edu\/\">Smithsonian<\/a> [exhibit: Written in Bone, How Bone Biographies Get Written] <a href=\"https:\/\/www.si.edu\/termsofuse\">is used for educational and non-commercial purposes as outlined by the Smithsonian.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Finally, <em>thermal trauma<\/em> is a bone alteration that results from bone exposure to extreme heat. Thermal trauma can result in cases of house or car fires, intentional disposal of a body in cases of homicidal violence, plane crashes, and so on. Thermal trauma is most often characterized by color changes to bone, ranging from yellow to black (charred) or white (calcined). Other bone alterations characteristic of thermal trauma include delamination (flaking or layering due to bone failure), shrinkage, fractures, and heat-specific burn patterning. When interpreting injuries resulting from thermal damage, an anthropologist can differentiate between thermal fractures and fractures that occurred before heat exposure, thereby contributing to the interpretation of burn patterning (e.g., was the individual bound or in a flexed position prior to the fire?).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">While there are characteristic patterns associated with the four categories of bone trauma, it is also important to note that these bone alterations do not always occur independently of different trauma types. An individual\u2019s skeleton may present with multiple different types of trauma, such as a projectile wound and thermal trauma. Therefore, it is important that the anthropologist recognize the different types of trauma and interpret them appropriately.<\/p>\n<h3 class=\"import-Normal\"><strong>Timing of Injury<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Another important component of any anthropological trauma analysis is the determination of the timing of injury (e.g., when did the injury occur). Timing of injury is traditionally split into one of three categories: <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1260\">antemortem<\/a> <\/strong>(before death), <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1258\">perimortem<\/a> <\/strong>(at or around the time of death), and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1266\">postmortem <\/a><\/strong>(after death). This classification system differs slightly from the classification system used by the pathologist because it specifically references the qualities of bone tissue and bone response to external forces. Therefore, the perimortem interval (at or around the time of death) means that the bone is still fresh and has what is referred to as a green bone response, which can extend past death by several weeks or even months. For example, in cold or freezing temperatures a body can be preserved for extended periods of time, increasing the perimortem interval, while in desert climates decomposition is accelerated, thereby significantly decreasing the postmortem interval (Galloway 1999, 12). Antemortem injuries (occurring well before death and not related to the death incident) are typically characterized by some level of healing, in the form of a fracture callus or unification of fracture margins. Finally, postmortem injuries (occurring after death, while bone is no longer fresh) are characterized by jagged fracture margins, resulting from a loss of moisture content during the decomposition process (Galloway 1999, 16). In general, all bone traumas should be classified according to the timing of injury, if possible. This information will help the medical examiner or pathologist better understand the circumstances surrounding the decedent\u2019s death, as well as events occurring during life and after the final disposition of the body.<\/p>\n<h3 class=\"import-Normal\"><strong>The Role of the Forensic Anthropologist in Trauma Analysis<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Within the medicolegal system, forensic anthropologists are often called upon by the medical examiner, forensic pathologist, or coroner to assist with an interpretation of trauma. The forensic anthropologist\u2019s main focus in any trauma analysis is the underlying skeletal system\u2014as well as, sometimes, cartilage. Analysis and interpretation of soft tissue injuries fall within the purview of the medical examiner or pathologist. It is also important to note that the main role of the forensic anthropologist is to provide information pertaining to skeletal injury to assist the medical examiner\/pathologist in their final interpretation of injury. Forensic anthropologists do not hypothesize as to the cause of death of an individual. Instead, a forensic anthropologist\u2019s report should include a description of the injury (e.g., trauma mechanism, number of injuries, location, timing of injury); documentation of the injury, which may be utilized in court testimony (e.g., photographs, radiographs, measurements); and, if applicable, a statement as to the condition of the body and state of decomposition, which may be useful for understanding the depositional context (e.g., how long has the body been exposed to the elements; was it moved or in its original location; are any of the alterations to bone due to environmental or faunal exposure instead of intentional human modification).<\/p>\n<h2 class=\"import-Normal\">Taphonomy<\/h2>\n<h2 class=\"import-Normal\"><strong>What Happened to the Remains After Death?<\/strong><\/h2>\n<p class=\"import-Normal\">The majority of the skeletal analysis process revolves around the identity of the deceased individual. However, there is one last, very important question that forensic anthropologists should ask: What happened to the remains after death? Generally speaking, processes that alter the bone after death are referred to as taphonomic changes (refer to Chapter 7 for a discussion regarding taphonomy and the fossil record).<\/p>\n<p class=\"import-Normal\">The term <em>taphonomy<\/em> was originally used to refer to the processes through which organic remains mineralize, also known as fossilization. Within the context of biological anthropology, the term <em>taphonomy<\/em> is better defined as the study of what happens to human remains after death (Komar and Buikstra 2008). Initial factors affecting a body after death include processes such as decomposition and scavenging by animals. However, taphonomic processes encompass much more than the initial period after death. For example, plant root growth can leach minerals from bone, leaving a distinctive mark. Sunlight can bleach human remains, leaving exposed areas whiter than those that remained buried. Water can wear the surface of the bone until it becomes smooth.<\/p>\n<p class=\"import-Normal\">Some taphonomic processes can help a forensic anthropologist estimate the relative amount of time that human remains have been exposed to the elements. For example, root growth through a bone would certainly indicate a body was buried for more than a few days. Forensic anthropologists must be very careful when attempting to estimate time since death based on taphonomic processes because environmental conditions can greatly influence the rate at which taphonomic processes progress. For example, in cold environments, tissue may decay slower than in warm, moist environments.<\/p>\n<p class=\"import-Normal\">Forensic anthropologists must contend with taphonomic processes that affect the preservation of bones. For example, high acidity in the soil can break down human bone to the point of crumbling. In addition, when noting trauma, they must be very careful not to confuse postmortem (after death) bone damage with trauma.<\/p>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 470.25pt\">\n<caption>Figure 15.21: Table showing taphonomic processes that affect the preservation of bones. A. Rodent gnawing. B. Carnivore damage. C. Burned bone. D. Root etching. E. Weathering. F. Cut marks. Credit: A. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Rodent gnawing (Figure 15.26)<\/a>, B. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Carnivore damage (Figure 15.27)<\/a>, C. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Burned bone (Figure 15.28)<\/a>, D. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Root etching (Figure 15.29)<\/a>, E. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Weathering (Figure 15.30)<\/a>, and F. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Cut marks (Figure 15.30)<\/a>, all original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone are under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-NC 4.0 License<\/a>.<\/caption>\n<thead>\n<tr style=\"height: 52.5pt\">\n<td class=\"Table1-C\" style=\"padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Taphonomic Process<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 1pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Definition<\/p>\n<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table1-R\" style=\"height: 190.5pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Rodent Gnawing<\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: center\"><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image19-2.png\" alt=\"Parallel tooth marks etched by a rodent\u2019s front teeth visible on the end of an animal bone.\" width=\"564\" height=\"422\" \/><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">When rodents, such as rats and mice, chew on bone, they leave sets of parallel grooves. The shallow grooves are etched by the rodent\u2019s incisors.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 166.75pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Carnivore Damage<\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: center\"><strong><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image23-4.png\" alt=\"Pit marks from the canines of a carnivore visible on the surface of an animal bone.\" width=\"410\" height=\"272\" \/><\/strong><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Carnivores may leave destructive dental marks on bone. The tooth marks may be visible as pit marks or punctures from the canines, as well as extensive gnawing or chewing of the ends of the bones to retrieve marrow.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 177pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Burned Bone<\/strong><\/p>\n<p class=\"import-Normal\"><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image20-5.png\" alt=\"Burned animal bone fragments pictured at different stages of thermal damage.\" width=\"512\" height=\"342\" \/><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Fire causes observable damage to bone. Temperature and the amount of time bone is heated affect the appearance of the bone. Very high temperatures can crack bone and result in white coloration. Color gradients are visible in between high and lower temperatures, with lower temperatures resulting in black coloration from charring. Cracking can also reveal information about the directionality of the burn.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 169.75pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Root Etching<\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: center\"><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image15-4.png\" alt=\"Animal bone with prominent, discolored grooves where roots leached nutrients from bone\u2019s surface.\" width=\"512\" height=\"342\" \/><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Plant roots can etch the outer surface of bone, leaving grooves where the roots attached as they leached nutrients. During this process, the plant\u2019s roots secrete acid that breaks down the surface of the bone.<\/p>\n<p class=\"import-Normal\">\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 170.5pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Weathering<\/strong><\/p>\n<p class=\"import-Normal\"><strong><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image9.png\" alt=\"Cracking and exfoliation of the surface of an animal bone. \" width=\"512\" height=\"342\" \/><\/strong><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Many different environmental conditions affect bone. River transport can smooth the surface of the bone due to water abrasion. Sunlight can bleach the exposed surface of bone. Dry and wet environments or the mixture of both types of environments can cause cracking and exfoliation of the surface. Burial in different types of soil can cause discoloration, and exposure can cause degreasing.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 169.75pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Cut Marks<\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: left\"><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image8-2.png\" alt=\"Thin vertical lines and cuts are visible along the bone.\" width=\"512\" height=\"342\" \/><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Humans may alter bone by cutting, scraping, or sawing it directly or in the process of removing tissue. The groove pattern\u2014that is, the depth and width of the cuts\u2014can help identify the tool used in the cutting process.<\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div class=\"textbox kk shaded\">\n<h2>Dig Deeper: Modern Forensic Technologies<\/h2>\n<p>In recent years, the forensics community has greatly benefited from the introduction of new technologies, helping strengthen the precision and speed of discoveries and advancements in the field. With recent developments in forensic anthropology, such as 3D scanning technologies, virtual reconstruction, and AI-assisted DNA analysis being integrated into traditional methods, there have been notable changes in how experts investigate human remains.<\/p>\n<p><strong>Artificial intelligence<\/strong><\/p>\n<p>In recent years, Artificial intelligence (AI) has shown itself to be a valuable tool within forensic anthropology. Aiding forensic experts and toxicologists with complex tasks, the limitations of traditional autopsies can be addressed with the help of AI. By automating and enhancing key investigative processes such as searching for microscopic changes in the human body to determine the cause of death or a person\u2019s life conditions, AI has the potential to enhance the efficiency of forensic processes significantly. It facilitates the detection of microscopic bodily changes to determine the cause of death or living conditions, compares evidence against databases for weapon identification and blood spatter analysis, and reduces manual workload. AI also enables the electronic storage of biometric data\u2013such as facial features, retinal patterns, and fingerprints\u2013for more accurate identity verification. Additionally, AI-powered microscopy enhances the detection of biological traces on complex surfaces, while blood biomarker analysis allows for more precise estimations of time of death (Wankhade et al., 2022).<\/p>\n<p>While AI holds great promise for the future of forensic medicine, a significant challenge remains: sourcing high-quality data to train the algorithms effectively. One of the more recent AI technologies making waves in the forensic anthropology sector is a new automated AI algorithm called the Convolutional Neural Network (CNN). As described by researchers in Switzerland\u2019s national medical journal Healthcare, CNN is a Deep Learning algorithm that allows for the detection of microscopic skull damage from CT scans or soft-tissue predictions of a face based on the skull information provided (Thurzo et al., 2021). While there are many advantages to using the CNN, the algorithm can be subject to biases in the same way human forensic experts can, as its assessment and pattern recognition of skulls and skeletons depend on the source data initially used for its AI training (2021).<\/p>\n<p><strong>3D Modeling<\/strong><\/p>\n<p>Identifying complex trauma to bones\u2013such as distinguishing heat fractures following blunt force trauma\u2013remains a significant challenge in forensic anthropology. This is particularly true for irregular skeletal structures like the pelvis, where overlapping trauma types can be difficult to differentiate, leading to these bones often being understudied. A 2024 study done by researchers from the University of Alberta in collaboration with the Michigan State Police explores the use of 3D laser scans and modelling technology to provide a highly detailed analysis of irregular bones with trauma. The study aimed to better distinguish peri-mortem trauma (trauma occurring around the time of death) from post-mortem heat alterations and improve the forensic analysis accuracy of such cases (Friedlander et al., 2024). The use of 3D laser scans and modelling technology provides very clear, detailed, and colored scans of bones, showing distinctions between the characteristics of the fractures. Blunt force and sharp force trauma produce a colour gradient on the 3D model that is more gradual and irregular, while heat fractures are more neat and characterized by little colour variation on the 3D models (2024). Other conclusions were also drawn from the study, such as the differences in trauma on fresh bones and bones that have been exposed to the elements for longer. An example of this is the interstitial fluid and collagen fibrils in fresh bones absorbing force, causing more long and jagged fracture lines, as opposed to a brittle fracture that older bones may exhibit (2024).<\/p>\n<p>Overall, the integration of 3D modeling technology offers a reproducible and highly detailed approach for analyzing trauma in anatomically complex and historically understudied skeletal regions. The practicality of this advancement is further emphasized by the researchers, who note that \u201cin many instances, scanned 3D models can be 3D printed for handheld representation of the model without damaging or overhandling the remains\u201d (2024, p. 2). By enhancing the ability to differentiate between various types of trauma and allowing for more convenient and risk-averse methods of research, this technology significantly improves the accuracy and reliability of forensic interpretations.<\/p>\n<\/div>\n<h2>Ethics and Human Rights<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Working with human remains requires a great deal of consideration and respect for the dead. Forensic anthropologists have to think about the ethics of our use of human remains for scientific purposes. How do we conduct casework in the most respectable manner possible? While there are a wide range of ethical considerations to consider when contemplating a career in forensic anthropology, this chapter will focus on two major categories: working with human remains and acting as an expert within the medicolegal system.<\/p>\n<h3 class=\"import-Normal\"><strong>Working with Human Remains<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropologists work with human remains in a number of contexts, including casework, excavation, research, and teaching. When working with human remains, it is always important to use proper handling techniques. To prevent damage to skeletal remains, bones should be handled over padded surfaces. Skulls should never be picked up by placing fingers in the eye orbits, foramen magnum (hole at the base of the skull for entry of the spinal cord), or through the zygomatic arches (cheekbones). Human remains, whether related to casework, fieldwork, donated skeletal collections, or research, were once living human beings. It is important to always bear in mind that work with remains should be ingrained with respect for the individual and their relatives. In addition to fieldwork, casework, and teaching, anthropologists are often invited to work with remains that come from a bioarchaeological context or from a human rights violation. While this discussion of ethics is not comprehensive, two case examples will be provided below in which an anthropologist must consider the ethical standards outlined above.<\/p>\n<h3 class=\"import-Normal\"><strong>Modern Human Rights Violations<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropologists may also be called to participate in criminal investigations involving human rights violations. Anthropological investigations may include assistance with identifications, determination of the number of victims, and trauma analyses. In this role, forensic anthropologists play an integral part in promoting human rights, preventing future human rights violations, and providing the evidence necessary to prosecute those responsible for past events. A few ethical considerations for the forensic anthropologist involved in human rights violations include the use of appropriate standards of identification, presenting reliable and unbiased testimony, and maintaining preservation of evidence. For a more comprehensive history of forensic anthropological contributions to human rights violations investigations, see Ubelaker 2018.<\/p>\n<h3 class=\"import-Normal\"><strong>Acting as an Expert in the Medicolegal System<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In addition to the ethical considerations involved in working with human skeletal remains, forensic anthropologists must abide by ethical standards when they act as experts within the medicolegal system. The role of the forensic anthropologist within the medicolegal system is primarily to provide information to the medical examiner or coroner that will aid in the identification process or determination of cause and manner of death. Forensic anthropologists also may be called to testify in a court of law. In this capacity, forensic anthropologists should always abide by a series of ethical guidelines that pertain to their interpretation, presentation, and preservation of evidence used in criminal investigations. First and foremost, practitioners should never misrepresent their training or education. When appropriate, outside opinions and assistance in casework should be requested (e.g., consulting a radiologist for radiological examinations or odontologist for dental exams). The best interest of the decedent should always take precedence. All casework should be conducted in an unbiased way, and financial compensation should never be accepted as it can act as an incentive to take a biased stance regarding casework. All anthropological findings should be kept confidential, and release of information is best done by the medical examiner or coroner. Finally, while upholding personal ethical standards, forensic anthropologists are also expected to report any perceived ethical violations committed by their peers.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ethical standards for the field of forensic anthropology are outlined by the Organization of Scientific Area Committees (OSAC) for Forensic Science, administered by the National Institute of Standards and Technology (NIST). OSAC and NIST recently began an initiative to develop standards that would strengthen the practice of forensic science both in the United States and internationally. OSAC\u2019s main objective is to \u201cstrengthen the nation\u2019s use of forensic science by facilitating the development of technically sound forensic science standards and by promoting the adoption of those standards by the forensic science community\u201d (NIST n.d.). Additionally, OSAC promotes the establishment of best practices and other guidelines to ensure that forensic science findings and their presentation are reliable and reproducible (NIST 2023).<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Native American Graves Protection and Repatriation Act (NAGPRA)<\/h2>\n<p class=\"import-Normal\">There is a long history in the <span style=\"background-color: #00ffff\">United States<\/span> of systematic disenfranchisement of Native American people, including lack of respect for tribal sovereignty. This includes the egregious treatment of Native American human remains. Over several centuries, thousands of Native American remains were removed from tribal lands and held at institutions in the United States, such as museums and universities.<\/p>\n<p class=\"import-Normal\">In 1990, a landmark human rights federal law, the Native American Graves Protection and Repatriation Act (NAGPRA), spurred change in the professional standards and practice of biological anthropology and archaeology. NAGPRA established a legal avenue to provide protection for and repatriation of Native American remains, cultural items, and sacred objects removed from Federal or tribal lands to Native American lineal descendants, Indian tribes, and Native Hawaiian organizations. Human remains and associated artifacts, curated in museum collections and federally funded institutions, are subject to three primary provisions outlined by the NAGPRA statute: (1) protection for Native graves on federal and private land; (2) recognition of tribal authority on such lands; and (3) the requirement that all Native skeletal remains and associated artifacts be inventoried and culturally affiliated groups be consulted concerning decisions related to ownership and final disposition (Rose, Green, and Green 1996). NAGPRA legislation was enacted to ensure ethical consideration and treatment of Native remains and to improve dialogue between scientists and Native groups.<\/p>\n<ul>\n<li>For more information about NAGPRA, visit the <a href=\"https:\/\/www.usbr.gov\/nagpra\/\" target=\"_blank\" rel=\"noopener\">Bureau of Reclamation NAGPRA website<\/a><\/li>\n<li>To read the text of the law, visit the <a href=\"https:\/\/www.congress.gov\/bill\/101st-congress\/house-bill\/5237\">US Congress NAGPRA law website<\/a>.<\/li>\n<li>For further discussion of NAGPRA history, please see <a href=\"https:\/\/textbooks.whatcom.edu\/tracesarchaeology\/\" target=\"_blank\" rel=\"noopener\"><em>TRACES: <\/em><em>An Open Invitation to <\/em><em>Archaeology <\/em>open textbook website<\/a><em><br \/>\n<\/em><\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Becoming a Forensic Anthropologist<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"background-color: #ff99cc\">What does it take to be a forensic anthropologist? Forensic anthropologists are first and foremost anthropologists. While many forensic anthropologists have an undergraduate degree in anthropology, they may also major in biology, criminal justice, pre-law, pre-med, and many other related fields. Practicing forensic anthropologists typically have an advanced degree, either a Master\u2019s or Doctoral degree in Anthropology. Additional training and experience in archaeology, the medico-legal system, rules of evidence, and expert witness testimony are also common. Practicing forensic anthropologists are also encouraged to be board-certified through the American Board of Forensic Anthropology (ABFA). Learn more about the field and educational opportunities on the ABFA website: <a class=\"rId111\" style=\"background-color: #ff99cc\" href=\"https:\/\/www.theabfa.org\/coursework\">https:\/\/www.theabfa.org\/coursework<\/a>.<\/span><\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Review Questions<\/h2>\n<ul>\n<li>What is forensic anthropology? What are the seven primary steps involved in a skeletal analysis?<\/li>\n<li>What are the major components of a biological profile? Why are forensic anthropologists often-tasked with creating biological profiles for unknown individuals?<\/li>\n<li>What are the four major types of skeletal trauma?<\/li>\n<li>What is taphonomy, and why is an understanding of taphonomy often critical in forensic anthropology analyses?<\/li>\n<li>What are some of the ethical considerations faced by forensic anthropologists?<\/li>\n<\/ul>\n<\/div>\n<h2>About the Authors<\/h2>\n<p><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image24-4.jpg\" alt=\"A woman with straight blonde hair smiles at the camera. \" width=\"191\" height=\"254\" \/><\/p>\n<h3 class=\"import-Normal\"><strong>Ashley Kendell, Ph.D.<\/strong><\/h3>\n<p class=\"import-Normal\">California State University, Chico, <a class=\"rId113\" href=\"mailto:akendell@csuchico.edu\">akendell@csuchico.edu<\/a><\/p>\n<p class=\"import-Normal\">Dr. Ashley Kendell is currently an associate professor and forensic anthropologist at Chico State. Prior to beginning her position at Chico State, she was a visiting professor at the University of Montana and the forensic anthropologist for the state of Montana. Dr. Kendell obtained her doctorate from Michigan State University, and her research interests include skeletal trauma analysis and digitization and curation methods for digital osteological data. She is also a Registry Diplomate of the American Board of Medicolegal Death Investigators. Throughout her doctoral program, she worked as a medicolegal death investigator for the greater Lansing, Michigan, area and was involved in the investigation of over 200 forensic cases.<\/p>\n<p class=\"import-Normal\"><strong><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image4.jpg\" alt=\"A woman with straight brown hair pulled back smiles at the camera. \" width=\"194\" height=\"258\" \/><\/strong><\/p>\n<h3 class=\"import-Normal\"><strong>Alex Perrone, M.A., M.S.N, R.N., P.H.N.<\/strong><\/h3>\n<p class=\"import-Normal\">Butte Community College, <a class=\"rId115\" href=\"mailto:perroneal@butte.edu\">perroneal@butte.edu<\/a><\/p>\n<p class=\"import-Normal\">Alex Perrone is a lecturer in anthropology at Butte Community College. She is also a Registered Nurse and a certified Public Health Nurse. She is a former Supervisor of the Human Identification Laboratory in the Department of Anthropology at California State University, Chico. Her research interests include bioarchaeology, paleopathology, forensic anthropology, skeletal biology, California prehistory, and public health. She has worked on bioarchaeological and archaeological projects in Antigua, California, Hawaii, Greece, and the UK, and was an archaeological technician for the USDA Forest Service. She assisted with training courses for local and federal law enforcement agencies and assisted law enforcement agencies with the recovery and analysis of human remains.<\/p>\n<p class=\"import-Normal\" data-wp-editing=\"1\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image1-1.jpg\" alt=\"A woman with curly brown, shoulder-length hair smiles at the camera.\" width=\"190\" height=\"253\" \/><\/p>\n<h3 class=\"import-Normal\"><strong>Colleen Milligan, Ph.D.<\/strong><\/h3>\n<p class=\"import-Normal\">California State University, Chico, <a class=\"rId117\" href=\"mailto:cfmilligan@csuchico.edu\">cfmilligan@csuchico.edu<\/a><\/p>\n<p class=\"import-Normal\">Dr. Colleen Milligan is a biological and forensic anthropologist with research interests in bioarchaeology, skeletal biology, and forensic anthropology. She has been a Fellow with the Department of Homeland Security and has assisted in forensic anthropology casework and recoveries in the State of Michigan and California. She has also assisted in community outreach programs in forensic anthropology and forensic science, as well as recovery training courses for local, state, and federal law enforcement officers. She is a certified instructor through Peace Officers Standards and Training (POST). Dr. Milligan serves as the current co-director of the Chico State Human Identification Laboratory.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration<\/h2>\n<p><a href=\"https:\/\/www.theabfa.org\/coursework\" target=\"_blank\" rel=\"noopener\">The American Board of Forensic Anthropology (ABFA)<\/a><\/p>\n<p><a href=\"https:\/\/www.aafs.org\/\" target=\"_blank\" rel=\"noopener\">The American Academy of Forensic Sciences (AAFS)<\/a><\/p>\n<p><a href=\"https:\/\/www.nist.gov\/organization-scientific-area-committees-forensic-science\" target=\"_blank\" rel=\"noopener\">The Organization of Scientific Area Committees for Forensic Science (OSAC)<\/a><\/p>\n<p><a href=\"https:\/\/textbooks.whatcom.edu\/tracesarchaeology\/\" target=\"_blank\" rel=\"noopener\">TRACES Bioarchaeology<\/a><\/p>\n<p><a href=\"https:\/\/transdoetaskforce.org\/\" target=\"_blank\" rel=\"noopener\">Trans Doe Task Force<\/a><\/p>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Adams, Bradley J., and Lyle W. Konigsberg, eds. 2008. <em>Recovery, Analysis, and Identification of Commingled Remains<\/em>. Totowa, NJ: Humana Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Beatrice, Jared S., and Angela Soler. 2016. \u201cSkeletal Indicators of Stress: A Component of the Biocultural Profile of Undocumented Migrants in Southern Arizona.\u201d <em>Journal of Forensic Sciences <\/em>61 (5): 1164\u20131172.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Berg, Gregory E. 2017. \u201cSex Estimation of Unknown Human Skeletal Remains.\u201d In <em>Forensic Anthropology: A Comprehensive Introduction, Second Edition<\/em>, edited by Natalie R. Langley and MariaTeresa A. Tersigni-Tarrant, 143\u2013159. Boca Raton, FL: CRC Press.<\/p>\n<p class=\"import-Normal\">Bethard, Jonathan D., and Elizabeth A. DiGangi. 2020. \u201cLetter to the Editor\u2014Moving Beyond a Lost Cause: Forensic Anthropology and Ancestry Estimates in the United States.\u201d <em>Journal of Forensic Sciences<\/em> 65 (5): 1791\u20131792.<\/p>\n<p class=\"import-Normal\">Birkby, Walter H., Todd W. Fenton, and Bruce E. Anderson. 2008. \u201cIdentifying Southwest Hispanics Using Nonmetric Traits and the Cultural Profile.\u201d <em>Journal of Forensic Sciences <\/em>53 (1): 29\u201333.<\/p>\n<p class=\"import-Normal\">Blatt, Samantha, Amy Michael, and Lisa Bright. Forthcoming. \u201cBioarchaeology: Interpreting Human Behavior from Skeletal Remains.\u201d In <em>TRACES: <\/em><em>An Open Invitation to <\/em><em>Archaeology<\/em>. https:\/\/textbooks.whatcom.edu\/tracesarchaeology\/.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Brooks, S., and J. M. Suchey. 1990. \u201cSkeletal Age Determination Based on the Os Pubis: A Comparison of the Acs\u00e1di-Nemesk\u00e9ri and Suchey-Brooks Methods.\u201d <em>Human Evolution <\/em>5 (3): 227\u2013238.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Buchanan, Shelby. 2014. \u201cBone Modification in Male to Female Transgender Surgeries: Considerations for the Forensic Anthropologist.\u201d MA thesis, Department of Geography and Anthropology, Louisiana State University, Baton Rouge.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Cunningham, Craig, Louise Scheuer, and Sue Black. 2016. <em>Developmental Juvenile Osteology, Second Edition<\/em>. London: Elsevier Academic Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Galloway, Alison, ed. 1999. <em>Broken Bones: Anthropological Analysis of Blunt Force Trauma<\/em>. Springfield, IL: Charles C. Thomas Publisher, LTD.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Hefner, Joseph T., and Kandus C. Linde. 2018. <em>Atlas of Human Cranial <\/em><em>Macromorphoscopic<\/em><em> Traits<\/em>. San Diego: Academic Press.<\/p>\n<p class=\"import-Normal\">\u0130\u015fcan, M. Y., S. R. Loth, and R. K. Wright. 1984. \u201cAge Estimation from the Rib by Phase Analysis: White Males.\u201d <em>Journal of Forensic Sciences <\/em>29 (4): 1094\u20131104.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">\u0130\u015fcan, M. Y., S. R. Loth, and R. K. Wright. 1985. \u201cAge Estimation from the Rib by Phase Analysis: White Females.\u201d <em>Journal of Forensic Sciences <\/em>30 (3): 853\u2013863.Katz, Darryl, and Judy Myers Suchey. 1986. \u201cAge Determination of the Male Os Pubis.\u201d <em>American Journal of Physical Anthropology <\/em>69 (4): 427\u2013435.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Komar, Debra A., and Jane E. Buikstra. 2008. <em>Forensic Anthropology: Contemporary Theory and Practice<\/em>. New York: Oxford University Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Langley, Natalie R., Alice F. Gooding, and MariaTeresa Tersigni-Tarrant. 2017. \u201cAge Estimation Methods.\u201d In <em>Forensic Anthropology: A Comprehensive Introduction, Second Edition<\/em>, edited by Natalie R. Langley and MariaTeresa A. Tersigni-Tarrant, 175\u2013191. Boca Raton, FL: CRC Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Lovell, Nancy C. 1997. \u201cTrauma Analysis in Paleopathology.\u201d <em>Yearbook of Physical Anthropology<\/em> 104 (S25): 139\u2013170.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Native American Graves Protection and Repatriation Act (NAGPRA) 1990 (25 U.S. Code 3001 et seq.)<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">NIST (National Institute of Standards and Technology). N.d. \u201cThe Organization of Scientific Area Committees for Forensic Science.\u201d Accessed April 18, 2023. <a class=\"rId120\" href=\"https:\/\/www.nist.gov\/topics\/organization-scientific-area-committees-forensic-science\">https:\/\/www.nist.gov\/topics\/organization-scientific-area-committees-forensic-science<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ousley, Stephen. 1995. \u201cShould We Estimate Biological or Forensic Stature?\u201d <em>Journal of Forensic Sciences<\/em> 40(5): 768\u2013773.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Phenice, T. W. 1969. \u201cA Newly Developed Visual Method of Sexing the Os Pubis.\u201d <em>American Journal of Physical Anthropology<\/em> 30 (2): 297\u2013302.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Rose, Jerome C., Thomas J. Green, and Victoria D. Green. 1996. \u201cNAGPRA Is Forever: Osteology and the Repatriation of Skeletons.\u201d <em>Annual Review of Anthropology <\/em>25: 81\u2013103.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Schaefer, Maureen, Sue Black, and Louise Scheuer. <em>Juvenile Osteology: A Laboratory and Field Manua<\/em>l. 2009. San Diego: Elsevier Academic Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Schall, Jenna L., Tracy L. Rogers, and Jordan D. Deschamps-Braly. 2020. \u201cBreaking the Binary: The Identification of Trans-women in Forensic Anthropology.\u201d <em>Forensic Science International<\/em> 309: 110220. https:\/\/doi.org\/10.1016\/j.forsciint.2020.110220.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2010a. \u201cPersonal Identification.\u201d Last modified June 3, 2010. <a class=\"rId121\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_personal_identification.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_personal_identification.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2010b. \u201cSex Assessment.\u201d Last modified June 3, 2010. <a class=\"rId122\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_sex_assessment.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_sex_assessment.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2011. \u201cTrauma Analysis.\u201d Last modified May 27, 2011. <a class=\"rId123\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_trauma.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_trauma.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2012. \u201cStature Estimation.\u201d Last modified August 2, 2012. <a class=\"rId124\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_stature_estimation.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_stature_estimation.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2013. \u201cAge Estimation.\u201d Last modified January 22, 2013. <a class=\"rId125\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_age_estimation.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_age_estimation.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Soler, Angela, and Jared S. Beatrice. 2018. \u201cExpanding the Role of Forensic Anthropology in Humanitarian Crisis: An Example from the USA-Mexico Border. In <em>Sociopolitics of Migrant Death and Repatriation: Perspectives from Forensic Science<\/em>, edited by Krista E. Latham and Alyson J. O\u2019Daniel, 115\u2013128. New York: Springer.<\/p>\n<p class=\"import-Normal\">Soler, Angela, Robin Reineke, Jared Beatrice, and Bruce E. Anderson. 2019. \u201cEtched in Bone: Embodied Suffering in the Remains of Undocumented Migrants.\u201d <em>In<\/em> <em>The Border and Its Bodies: The Embodiment of Risk along the U.S.-M\u00e9xico Line<\/em>, edited by Thomas E. Sheridan and Randall H. McGuire, 173\u2013207. Tucson: University of Arizona Press.<\/p>\n<p class=\"import-Normal\">Stull, Kyra E., Eric J. Bartelink, Alexandra R. Klales, Gregory E. Berg, Michael W. Kenyhercz, Erica N. L\u2019Abb\u00e9, Matthew C. Go, et al.. 2021. \u201cCommentary on: Bethard JD, DiGangi EA. Letter to the Editor\u2014Moving Beyond a Lost Cause: Forensic Anthropology and Ancestry Estimates in the United States. J Forensic Sci. 2020;65(5):1791\u20132. doi: 10.1111\/1556-4029.14513.\u201d <em>Journal of Forensic Sciences <\/em>66 (1): 417\u2013420.<\/p>\n<p class=\"import-Normal\">Tallman, Sean D., Caroline D. Kincer, and Eric D. Plemons. 2022. \u201cCentering Transgender Individuals in Forensic Anthropology and Expanding Binary Sex Estimation in Casework and Research.\u201d Special issue, \u201cDiversity and Inclusion,\u201d <em>Forensic Anthropology<\/em> 5 (2): 161\u2013180.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Tersigni-Tarrant, MariaTeresa A., and Natalie R. Langley. 2017. \u201cHuman Osteology.\u201d In <em>Forensic Anthropology: A Comprehensive Introduction, Second Edition<\/em>, edited by Natalie R. Langley and MariaTeresa A. Tersigni-Tarrant, 81\u2013109. Boca Raton, FL: CRC Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ubelaker, Douglas H. 2018. \u201cA History of Forensic Anthropology.\u201d Special issue, \u201cCentennial Anniversary Issue of AJPA,\u201d <em>American Journal of Physical Anthropology<\/em> 165 (4): 915\u2013923.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">White, Tim D., and Pieter A. Folkens. 2005. <em>The Human Bone Manual<\/em>. Burlington, MA: Elsevier Academic Press.<\/p>\n<p class=\"import-Normal\">Winburn, Allysha P., and Bridget Algee-Hewitt. 2021. \u201cEvaluating Population Affinity Estimates in Forensic Anthropology: Insights from the Forensic Anthropology Database for Assessing Methods Accuracy (FADAMA).\u201d <em>Journal of Forensic Sciences<\/em> 66 (4): 1210\u20131219.<\/p>\n<p class=\"import-Normal\">Winburn, Allysha Powanda, Sarah Kiley Schoff, and Michael W. Warren. 2016. \u201cAssemblages of the Dead: Interpreting the Biocultural and Taphonomic Signature of Afro- Cuban Palo Practice in Florida.\u201d <em>Journal of African Diaspora Archaeology and Heritage <\/em>5 (1): 1\u201337.<\/p>\n<\/div>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1783\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1783\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\">Ashley Kendell, Ph.D., California State University, Chico<\/p>\n<p class=\"import-Normal\">Alex Perrone, M.A., M.S.N, R.N., P.H.N., Butte Community College<\/p>\n<p class=\"import-Normal\">Colleen Milligan, Ph.D., California State University, Chico<\/p>\n<p class=\"import-Normal\"><em>This chapter is a revision from \"<\/em><a class=\"rId7\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\"><em>Chapter 15: Bioarchaeology and Forensic Anthropology<\/em><\/a><em>\u201d by Ashley Kendell, Alex Peronne, and Colleen Milligan. In <\/em><a class=\"rId8\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, first edition<\/em><\/a><em>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under <\/em><a class=\"rId9\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\"><em>CC BY-NC 4.0<\/em><\/a><em>. <\/em><\/p>\n<p class=\"import-Normal\"><strong>Content Warning and Disclaimer:<\/strong> This chapter includes images of human remains as well as discussions centered on human skeletal analyses. All images are derived from casts, sketches, nonhuman skeletal material, as well as non-Indigenous skeletal materials curated within the CSU, Chico Human Identification Lab, and the Hartnett-Fulginiti donated skeletal collection.<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\"><span style=\"color: #ffffff\">Learning Objectives<\/span><\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Define forensic anthropology as a subfield of biological anthropology.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Describe the seven steps carried out during skeletal analysis.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Outline the four major components of the biological profile.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Contrast the four categories of trauma.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Explain how to identify the different taphonomic agents that alter bone.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 18pt\">Discuss ethical considerations for forensic anthropology.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1214\">Forensic anthropology<\/a><\/strong> is a subfield of biological anthropology and an applied area of anthropology. Forensic anthropologists use skeletal analysis to gain information about humans in the present or recent past, then they apply this information within a medicolegal context. This means that forensic anthropologists specifically conduct their analysis on recently deceased individuals (typically within the last 50 years) as part of investigations by law enforcement. Forensic anthropologists can assist law enforcement agencies in several different ways, including aiding in the identification of human remains whether they are complete, fragmentary, burned, scattered, or decomposed. Additionally, forensic anthropologists can help determine what happened to the deceased at or around the time of death as well as what processes acted on the body after death (e.g., whether the remains were scattered by animals, whether they were buried in the ground, or whether they remained on the surface as the soft tissue decomposed).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Many times, because of their expertise in identifying human skeletal remains, forensic anthropologists are called to help with outdoor search-and-recovery efforts, such as locating remains scattered across the surface or carefully excavating and documenting buried remains. In other cases, forensic anthropologists recover remains after natural disasters or accidents, such as fire scenes, and can help identify whether each bone belongs to a human or an animal. Forensic anthropology spans a wide scope of contexts involving the law, including incidences of mass disasters, genocide, and war crimes.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">A point that can be somewhat confusing for students is that although the term <em>forensic<\/em> is included in this subfield of biological anthropology, there are many forensic techniques that are not included in the subfield. Almost exclusively, forensic anthropology deals with skeletal analysis. While this can include the comparison of antemortem (before death) and postmortem (after death) radiographs to identify whether remains belong to a specific person, or using photographic superimposition of the cranium, it does not include analyses beyond the skeleton. For example, blood-spatter analysis, DNA analysis, fingerprints, and material evidence collection do not fall under the scope of forensic anthropology.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">So, what can forensic anthropologists glean from bones alone? Forensic anthropologists can address a number of questions about a human individual based on their skeletal remains. Some of those questions are as follows: How old was the person? Was the person biologically male or female? How tall was the person? What happened to the person at or around their time of death? Were they sick? The information from the skeletal analysis can then be matched with missing persons records, medical records, or dental records, aiding law enforcement agencies with identifications and investigations.<\/p>\n<h2 class=\"import-Normal\">Skeletal Analysis<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropology relies on skeletal analysis to reveal information about the deceased. <span style=\"background-color: #00ffff\">The methodology and approaches outlined below are specific to the United States.<\/span> Forensic anthropological methods differ depending on the country conducting an investigation. In the United States, there are typically seven steps or questions to the process:<\/p>\n<ul>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Is it bone?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Is it human?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Is it modern or archeological?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">How many individuals are present or what is the minimum number of individuals (MNI)?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Who is it?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Is there evidence of trauma before or around the time of death?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">What happened to the remains after death?<\/li>\n<\/ul>\n<h3 class=\"import-Normal\"><strong>Is It Bone?<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">One of the most important steps in any skeletal analysis starts with determining whether or not material suspected to be bone is in fact bone. Though it goes without saying that a forensic anthropologist would only carry out analysis on bone, this step is not always straightforward. Whole bones are relatively easy to identify, but determining whether or not something is bone becomes more challenging once it becomes fragmentary. As an example, in high heat such as that seen on fire scenes, bone can break into pieces. During a house fire with fatalities, firefighters watered down the burning home. After the fire was extinguished, the sheetrock (used to construct the walls of the home) was drenched and crumbled. The crumbled sheetrock was similar in color and form to burned, fragmented bone, therefore mistakable for human remains (Figure 15.1). Forensic anthropologists on scene were able to separate the bones from the construction material, helping to confirm the presence of bone and hence the presence of individual victims of the fire. In this case, forensic anthropologists were able to recognize the anatomical and layered structure of bone and were able to distinguish it from the uniform and unlayered structure of sheetrock.<\/p>\n<p class=\"import-Normal\"><strong><img class=\"aligncenter\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/06\/image3.png\" alt=\"Long rectangular sheetrock with exposed porous surface.\" width=\"182\" height=\"208\" \/><\/strong><\/p>\n<figure style=\"width: 372px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image6-1.png\" alt=\"Two examples of sheetrock with dried or burnt surfaces.\" width=\"372\" height=\"210\" \/><figcaption class=\"wp-caption-text\">Figure 15.1: Burned sheetrock used as building material appears similar to human bone but can be differentiated by the fact that it is the same density throughout. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Example of burned sheetrock (Figure 15.1)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">As demonstrated by the example above, both the macrostructure (visible with the naked eye) and microstructure (visible with a microscope) of bone are helpful in bone identification. Bones are organs in the body made up of connective tissue. The connective tissue is hardened by a mineral deposition, which is why bone is rigid in comparison to other connective tissues such as cartilage (Tersigni-Tarrant and Langley 2017, 82\u201383; White and Folkens 2005, 31). In a living body, the mineralized tissue does not make up the only component of bone\u2014there are also blood, bone marrow, cartilage, and other types of tissues. However, in dry bone, two distinct layers of the bone are the most helpful for identification. The outer layer is made up of densely arranged osseous (bone) tissue called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1216\">compact (cortical) bone<\/a><\/strong>. The inner layer is composed of much more loosely organized, porous bone tissue whose appearance resembles that of a sponge, hence the name <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1218\"><strong>spongy (trabecular) bone<\/strong><\/a>. Knowing that most bone contains both layers helps with the macroscopic identification of bone (Figures 15.2, 15.3). For example, a piece of coconut shell might look a lot like a fragment of a human skull bone. However, closer inspection will demonstrate that coconut shell only has one very dense layer, while bone has both the compact and spongy layers.<\/p>\n<figure style=\"width: 380px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image27-1.png\" alt=\"Drawing showing thick exterior compact bone and porous internal cortical bone.\" width=\"380\" height=\"371\" \/><figcaption class=\"wp-caption-text\">Figure 15.2: Cross section of human long bone with compact and cortical bone layers visible. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Cross section of human long bone (Figure 15.2)<\/a> original to<a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"> Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p>&nbsp;<\/p>\n<figure style=\"width: 364px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image25-2.png\" alt=\"Cranial bone cross section called a periosteum with spongy bone (diploe) and compact bone labeled. Compact bone is a thin slice at the top and bottom and is smooth and hard. Spongy bone is in the middle and has irregular holes and indentations throughout. \" width=\"364\" height=\"184\" \/><figcaption class=\"wp-caption-text\">Figure 15.3: Cranial anatomy is slightly different as compared to that of a long bone in cross section. The compact (cortical) bone layers sandwich the spongy (trabecular) bone. One layer of compact bone forms the very outer surface of the skull and the other lines the internal surface of the skull. Credit: <a href=\"https:\/\/cnx.org\/contents\/FPtK1zmh@6.27:kwbeYj9S@3\/Bone-Structure\">Anatomy of a Flat Bone (Anatomy &amp; Physiology, Figure 6.3.3)<\/a> by<a href=\"https:\/\/openstax.org\/\"> OpenStax<\/a> is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/\"> CC BY 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The microscopic identification of bone relies on knowledge of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1220\">osteons<\/a><\/strong>, or bone cells (Figure 15.4). Under magnification, bone cells are visible in the outer, compact layer of bone. The bone cells are arranged in a concentric pattern around blood vessels for blood supply. The specific shape of the cells can help differentiate, for example, a small piece of PVC (white plastic) pipe from a human bone fragment (Figure 15.5).<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 340px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image14-3.png\" alt=\"Microscope image showing clustered osteons. Each has many rings and a dark center.\" width=\"340\" height=\"218\" \/><figcaption class=\"wp-caption-text\">Figure 15.4: Bone microstructure (osteons). Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Bone_(248_12)_Bone_cross_section.jpg\">Bone (248 12) Bone cross section<\/a> by <a href=\"https:\/\/cs.wikipedia.org\/wiki\/Josef_Reischig\">Doc. RNDr. Josef Reischig, CSc.<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p>&nbsp;<\/p>\n<figure style=\"width: 332px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image7-1.png\" alt=\"Flat, white section of PVC. Edges are broken and surface rough.\" width=\"332\" height=\"268\" \/><figcaption class=\"wp-caption-text\">Figure 15.5: Fragments of plastic PVC pipe, such as those seen in this photo, may be mistaken for human bone. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Example of PVC pipe<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Is It Human?<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Once it has been determined that an object is bone, the next logical step is to identify whether the bone belongs to a human or an animal. Forensic anthropologists are faced with this question in everyday practice because human versus nonhuman bone identification is one of the most frequent requests they receive from law enforcement agencies.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">There are many different ways to distinguish human versus nonhuman bone. The morphology (the shape\/form) of human bone is a good place for students to start. Identifying the 206 bones in the adult human skeleton and each bone\u2019s distinguishing features (muscle attachment sites, openings and grooves for nerves and blood vessels, etc.) is fundamental to skeletal analysis.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Nevertheless, there are many animal bones and human bones that look similar. For example, the declawed skeleton of a bear paw looks a lot like a human hand, pig molars appear similar to human molars, and some smaller animal bones might be mistaken for those of an infant. To add to the confusion, fragmentary bone may be even more difficult to identify as human or nonhuman. However, several major differences between human and nonhuman vertebrate bone help distinguish the two.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropologists pay special attention to the density of the outer, compact layer of bone in both the cranium and in the long bones. Human cranial bone has three distinctive layers. The spongy bone is sandwiched between the outer (ectocranial) and inner (endocranial) compact layers. In most other mammals, the distinction between the spongy and compact layers is not always so definite. Secondly, the compact layer in nonhuman mammal long bones can be much thicker than observed in human bone. Due to the increased density of the compact layer, nonhuman bone tends to be heavier than human bone (Figure 15.6).<\/p>\n<figure style=\"width: 399px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image11-2.png\" alt=\"Ring-like cross section of bone.\" width=\"399\" height=\"266\" \/><figcaption class=\"wp-caption-text\">Figure 15.6: The compact layer of this animal bone is very thick, with almost no spongy bone visible. Compare with Figure 15.2 to visualize the difference in structure between human and nonhuman bone. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Animal bone cross section (Figure 15.6)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The size of a bone can also help determine whether it belongs to a human. Adult human bones are larger than subadult or infant bones. However, another major difference between human adult bones and those of a young individual or infant human can be attributed to development and growth of the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1222\">epiphyses<\/a><\/strong> (ends of the bone). The epiphyses of human subadult bones are not fused to the shaft (Figure 15.7). Therefore, if a bone is small and it is suspected to belong to a human subadult or infant, the epiphyses would not be fused. Many small animal bones appear very similar in form compared to adult human bones, but they are much too small to belong to an adult human. Yet they can be eliminated as subadult or infant bones if the epiphyses are fused to the shaft.<\/p>\n<figure style=\"width: 288px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image13-3.png\" alt=\"X-ray image of child\u2019s ankle.\" width=\"288\" height=\"412\" \/><figcaption class=\"wp-caption-text\">Figure 15.7: An x-ray of a subadult\u2019s ankle with the epiphyses of the tibia and fibula visible. The gap between the shaft of the bone and the end of the bone (epiphysis) is the location of the growth plate. Therefore, the growth plate gap is what separates the shafts from the epiphyses in the image. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Tib_fib_growth_plates.jpg\">Tib fib growth plates<\/a> by <a href=\"https:\/\/en.wikipedia.org\/wiki\/User:Gilo1969\">Gilo1969<\/a> at <a href=\"https:\/\/en.wikipedia.org\/wiki\/\">English Wikipedia<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/3.0\/legalcode\">CC BY 3.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Is It Modern or Archaeological? <\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropologists work with modern cases that fall within the scope of law enforcement investigations. Accordingly, it is important to determine whether discovered human remains are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1224\">archaeological<\/a> <\/strong>or forensic in nature. Human remains that are historic are considered archeaological. The scientific study of human remains from archaeological sites is called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1226\">bioarchaeology<\/a><\/strong>.<\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Dig Deeper: Bioarchaeology<\/h2>\n<p class=\"import-Normal\">For readers who are interested in the sister subfield of bioarchaeology, which studies human remains and material culture from the past, please refer to chapter 8 of <em>Bioarchaeology: Interpreting Human Behavior from Skeletal Remains,<\/em> in <em>TRACES: An Open Invitation to Archaeology<\/em> (Blatt, Michael, and Bright forthcoming).<\/p>\n<\/div>\n<p>A forensic anthropologist should begin their analysis by reviewing the context in which the remains were discovered. This will help them understand a great deal about the remains, including determining whether they are archaeological or forensic in nature as well as considering legal and ethical issues associated with the collection, analysis, and storage of human remains (see \u201cEthics and Human Rights\u201d section of this chapter for more information).<\/p>\n<figure style=\"width: 403px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image10-3.png\" alt=\"Four teeth in a person\u2019s mouth. First molar with silver filling.\" width=\"403\" height=\"303\" \/><figcaption class=\"wp-caption-text\">Figure 15.8: A human tooth with a filling. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Filling.jpg#filehistory\">Filling<\/a> by Kauzio has been designated to the <a href=\"https:\/\/creativecommons.org\/share-your-work\/public-domain\/cc0\/\">public domain (CC0)<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The \u201ccontext\u201d refers to the relationship the remains have to the immediate area in which they were found. This includes the specific place where the remains were found, the soil or other organic matter immediately surrounding the remains, and any other objects or artifacts in close proximity to the body. For example, imagine that a set of remains has been located during a house renovation. The remains are discovered below the foundation. Do the remains belong to a murder victim? Or was the house built on top of an ancient burial ground? Observing information from the surroundings can help determine whether the remains are archaeological or modern. How long ago was the foundation of the house erected? Are there artifacts in close proximity to the body, such as clothing or stone tools? These are questions about the surroundings that will help determine the relative age of the remains.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Clues directly from the skeleton may also indicate whether the remains are archaeological or modern. For example, tooth fillings can suggest that the individual was alive recently (Figure 15.8). In fact, filling material has changed over the decades, so the specific type of material used to fix a cavity can be matched with specific time periods. Gold was used in dental work in the past, but more recently composite (a mixture of plastic and fine glass) fillings have become more common.<\/p>\n<h3><strong>How <\/strong><strong>Many Individuals Are Present?<\/strong><\/h3>\n<h4 class=\"import-Normal\"><em>What Is MNI?<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Another assessment that an anthropologist can perform is the calculation of the number of individuals in a mixed burial assemblage. Because not all burials consist of a single individual, it is important to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1268\">burial assemblage<\/a><\/strong> be able to estimate the number of individuals in a forensic context. Quantification of the number of individuals in a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1524\">burial assemblage<\/a><\/strong> can be done through the application of a number of methods, including the following: the Minimum Number of Individuals (MNI), the Most Likely Number of Individuals (MLNI), and the Lincoln Index (LI). The most commonly used method in biological anthropology, and the focus of this section, is determination of the MNI.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The MNI presents \u201cthe minimum estimate for the number of individuals that contributed to the sample\u201d (Adams and Konigsberg 2008, 243). Many methods of calculating MNI were originally developed within the field of zooarchaeology for use on calculating the number of individuals in faunal or animal assemblages (Adams and Konigsberg 2008, 241). What MNI calculations provide is a lowest possible count for the total number of individuals contributing to a skeletal assemblage. Traditional methods of calculating MNI include separating a skeletal assemblage into categories according to the individual bone and the side the bone comes from and then taking the highest count per category and assigning that as the minimum number (Figure 15.9).<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 664px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image28-3.png\" alt=\"Many bone portions laying on individual plastic bags on a table.\" width=\"664\" height=\"441\" \/><figcaption class=\"wp-caption-text\">Figure 15.9: Skeletal elements from a commingled faunal assemblage. Credit: Commingled animal remains from Eden-Farson Pre-Contact site in southwest Wyoming by Matt O\u2019Brien original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Why Calculate MNI?<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In a forensic context, the determination of MNI is most applicable in cases of mass graves, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1232\">commingled burials<\/a><\/strong>, and mass fatality incidents. The term <em>commingled<\/em> is applied to any burial assemblage in which individual skeletons are not separated into separate burials. As an example, the authors of this chapter have observed commingling of remains resulting from mass fatality wildfire events. Commingled remains may also be encountered in events such as a plane or vehicle crash. It is important to remember that in any forensic context, MNI should be referenced and an MNI of one should be substantiated by the fact that there was no repetition of elements associated with the case.<\/p>\n<h3 class=\"import-Normal\"><strong>Constructing the Biological Profile<\/strong><\/h3>\n<h4 class=\"import-Normal\"><em>Who Is It?<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">\u201cWho is it?\u201d is one of the first questions that law enforcement officers ask when they are faced with a set of skeletal remains. To answer this question, forensic anthropologists construct a biological profile (White and Folkens 2005, 405). A <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1228\">biological profile<\/a> <\/strong>is an individual\u2019s identifying characteristics, or biological information, which include the following: biological sex, age at death, stature, population affinity, skeletal variation, and evidence of trauma and pathology.<\/p>\n<h4 class=\"import-Normal\"><em>Assessing Biological Sex <\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Assessment of biological sex is often one of the first things considered when establishing a biological profile because several other parts, such as age and stature estimations, rely on an assessment of biological sex to make the calculations more accurate.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Assessment of biological sex focuses on differences in both morphological (form or structure) and metric (measured) traits in individuals. When assessing morphological traits, the skull and the pelvis are the most commonly referenced areas of the skeleton. These differences are related to sexual dimorphism usually varying in the amount of robusticity seen between males and females. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1230\">Robusticity<\/a> <\/strong>deals with strength and size; it is frequently used as a term to describe a large size or thickness. In general, males will show a greater degree of robusticity than females. For example, the length and width of the mastoid process, a bony projection located behind the opening for the ear, is typically larger in males. The mastoid process is an attachment point for muscles of the neck, and this bony projection tends to be wider and longer in males. In general, cranial features tend to be more robust in males (Figure 15.10).<\/p>\n<figure style=\"width: 601px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image26-3.png\" alt=\"Front and side images of a male (left) and female (right) cranium.\" width=\"601\" height=\"632\" \/><figcaption class=\"wp-caption-text\">Figure 15.10: Anterior and lateral view of a male and female cranium. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Anterior and lateral view of a male and female cranium (Figure 15.10)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropo logy<\/a> by Ashley Kendell is a collective work under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Includes <a href=\"https:\/\/boneclones.com\/product\/modern-human-asian-female-skull-BC-149\/category\/all-human-skulls\/human-anatomy\">Human Female Asian Skull<\/a> and <a href=\"https:\/\/boneclones.com\/product\/human-asian-male-skull-BC-016\/category\/all-human-skulls\/human-anatomy\">Human Male Asian Skull<\/a> by <a href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a>, used by permission.]<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">When considering the pelvis, the features associated with the ability to give birth help distinguish females from males. During puberty, estrogen causes a widening of the female pelvis to allow for the passage of a baby. Several studies have identified specific features or bony landmarks associated with the widening of the hips, and this section will discuss one such method. The Phenice Method (Phenice 1969) is traditionally the most common reference used to assess morphological characteristics associated with sex. The Phenice Method specifically looks at the presence or absence of (1) a ventral arc, (2) the presence or absence of a subpubic concavity, and (3) the width of the medial aspect of the ischiopubic ramus (Figure 15.11). When present, the ventral arc, a ridge of bone located on the ventral surface of the pubic bone, is indicative of female remains. Likewise the presence of a subpubic concavity and a narrow medial aspect of the ischiopubic ramus is associated with a female sex estimation. Assessments of these features, as well as those of the skull (when both the pelvis and skull are present), are combined for an overall estimation of sex.<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 1603px\" class=\"wp-caption alignnone\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image29-3.png\" alt=\"Male and female os coxae (anterior portions).\" width=\"1603\" height=\"582\" \/><figcaption class=\"wp-caption-text\">Figure 15.11: Features associated with the Phenice Method. Images derived from CSU-HIL donated skeletal collection. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Features associated with the Phenice Method (Figure 15.11)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Colleen Milligan is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Metric analyses are also used in the estimation of sex. Measurements taken from every region of the body can contribute to estimating sex through statistical approaches that assign a predictive value of sex. These approaches can include multiple measurements from several skeletal elements in what is called multivariate (multiple variables) statistics. Other approaches consider a single measurement, such as the diameter of the head of the femur, of a specific element in a univariate (single variable) analysis (Berg 2017, 152\u2013156).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">It is important to note that, although forensic anthropologists usually begin assessment of biological profile with biological sex, there is one major instance in which this is not appropriate. The case of two individuals found in California, on July 8, 1979, is one example that demonstrates the effect age has on the estimation of sex. The identities of the two individuals were unknown; therefore, law enforcement sent them to a lab for identification. A skeletal analysis determined that the remains represented one adolescent male and one adolescent female, both younger than 18 years of age. This information did not match with any known missing children at the time.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In 2015, the cold case was reanalyzed, and DNA samples were extracted. The results indicated that the remains were actually those of two girls who went missing in 1978. The girls were 15 years old and 14 years old at the time of death. It is clear that the 1979 results were incorrect, but this mistake also provides the opportunity to discuss the limitations of assessing sex from a subadult skeleton.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Assessing sex from the human skeleton is based on biological and genetic traits associated with females and males. These traits are linked to differences in sexual dimorphism and reproductive characteristics between females and males. The link to reproductive characteristics means that most indicators of biological sex do not fully manifest in prepubescent individuals, making estimations of sex unreliable in younger individuals (SWGANTH 2010b). This was the case in the example of the 14-year-old girl. When examined in 1979, her remains were misidentified as male because she had not yet fully developed female pelvic traits.<\/p>\n<h4 class=\"import-Normal\"><em>Sex vs. Gender<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Biological sex is a different concept than <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1234\">gender<\/a><\/strong>. While biological anthropologists can estimate sex from the skeleton, estimating an individual\u2019s gender would require a greater context because gender is defined culturally rather than biologically. Take, for example, an individual who identifies as transgender. This individual has a gender identity that is different from their biological sex. The gender identity of any individual depends on factors related to self-identification, situation or context, and cultural factors. <span style=\"background-color: #00ffff\">While in the U.S<\/span>. we have historically thought of sex and gender as binary concepts (male or female), many cultures throughout the world recognize several possible gender identities. In this sense, gender is seen as a continuous or fluid variable rather than a fixed one.<\/p>\n<p class=\"import-Normal\">Historically, forensic anthropologists have used a binary construct to categorize human skeletal remains as either male or female (with the accompanying categories of probable male, probable female, and indeterminate). In the case of transgender and gender nonconforming individuals, the binary approach to sex assessment may delay or hinder identification efforts (Buchanan 2014; Schall, Rogers, and Deschamps-Braly 2020; Tallman, Kincer, and Plemons 2021). As such, many forensic anthropologists have begun to address the inherent problems associated with a binary approach to sex identification and to explore ways of assessing social identity and self-identified gender using skeletal remains and forensic context.<\/p>\n<p class=\"import-Normal\">For the duration of this section, the term <em>transgender<\/em> refers to individuals whose gender identity differs from the sex assigned at birth (Schall, Rogers, and Deschamps-Braly 2020:2). Transgender individuals transition from one gender binary to another, such as male-to-female (MTF) or female-to-male (FTM). While many of the gender-affirming procedures available to trans and gender-nonconforming individuals are focused on soft tissue modifications (e.g., breast augmentation, genital reconstruction, hormone therapies, etc.), there are a number of gender-affirmation surgeries that do leave a permanent record on the skeleton. Generally speaking, FTM transgender people are reported to undergo fewer surgical procedures than do MTF transgender people (Buchanan 2014). The discussion below focuses on Facial Feminization Surgery (FFS), which leaves a permanent record on the human skeleton that may be used to help make an identification.<\/p>\n<p class=\"import-Normal\">FFS refers to a combination of procedures focused on sexually dimorphic features of the face, with the intent of transforming typically male facial features into more feminine forms. Facial Feminization Surgery procedures were developed by Dr. Douglas Ousterhout, a San Francisco based cranio-maxillofacial surgeon, in the mid-1980s (Schall, Rogers, and Deschamps-Braly 2020:2). FFS can include one or a combination of the following: hairline lowering, forehead reduction and contouring, brow lift, reduction rhinoplasty, cheek enhancement, lift lift, lip filling, chin contouring, jaw contouring, and\/or tracheal shave (Buchanan 2014; Schall, Rogers, and Deschamps-Braly 2020:2). Of the procedures outlined previously, four are known to directly affect the facial skeleton: forehead contouring, rhinoplasty, chin contouring, and jaw contouring (Buchanan 2014; Schall, Rogers, and Deschamps-Braly 2020:2).<\/p>\n<p class=\"import-Normal\">Because FFS procedures have been widely documented in the medical (and more recently the forensic anthropological) literature, there are a number of indicators that a forensic anthropologist can use to make more informed evaluations of gender, including evidence of bone remodeling in sexually dimorphic regions of the skull (e.g., forehead, chin, jawline), as well as the presence of plates, pins, or other surgical hardware that may be evidence of FFS (Buchanan 2014; Schall, Rogers, and Deschamps-Braly 2020; Tallman, Kincer, and Plemons 2021). Additionally, some forensic anthropologists suggest cautiously integrating contextual information from the scene, such as personal effects, material evidence, and recovery scene information, into their evaluation of an individual\u2019s social identity (Beatrice and Soler 2016; Birkby, Fenton, and Anderson 2008; Soler and Beatrice 2018; Soler et al. 2019; Tallman, Kincer, and Plemons 2021; Winburn, Schoff, and Warren 2016). The ultimate goal of many skeletal analyses is to make a positive identification on a set of unidentified remains.<\/p>\n<h4 class=\"import-Normal\"><em>Assessment <\/em><em>of Population Affinity<\/em><\/h4>\n<p>In an effort to combat the erroneous assumptions tied to the race concept, forensic anthropologists have attempted to reframe this component of the biological profile. The term <em>race<\/em> is no longer used in casework and teaching. Historically, the word <em>ancestry<\/em> is and was deemed a more appropriate way to describe an individual\u2019s phenotype. However, in more recent years, forensic anthropologists have begun using the term <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1236\">population affinity<\/a><\/strong><em>, <\/em>recognizing that we are basing our analysis on the similarities we see based on the reference samples we have available (Winburn and Algee-Hewitt 2021). An important note here is that it is possible to hinder identifications and harm individuals when tools like estimations of population affinity are misapplied, misinterpreted, or misused. For this reason, the field of forensic anthropology has ongoing conversations about the appropriateness of this analysis in the biological profile (Bethard and DiGangi 2020; Stull et al. 2021).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">We use the term <em>population affinity<\/em> to refer to the variation seen among modern populations\u2014variation that is both genetic and environmentally driven. The word <em>affinity<\/em> refers to similarities or relationships between individuals. As forensic anthropologists, we compare an unknown individual to multiple reference groups and look for the degree of similarity in observable traits with those groups. As noted previously, population affinity can aid law enforcement in their identification of missing persons or unknown skeletal remains.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Within the field of anthropology, the estimation of population affinity has a contentious history, and early attempts at classification were largely based on the erroneous assumption that an individual\u2019s <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1238\">phenotype <\/a><\/strong> (outward appearance) was correlated with their innate intelligence and abilities (see Chapter 13 for a more in-depth discussion of the history of the race concept). The use of the term <em>race<\/em> is deeply embedded in the social context of the United States. In any other organism\/living thing, groups divided according to the biological race concept would be defined as a separate subspecies. The major issue with applying the biological race concept to humans is that there are not enough differences between any two populations to separate on a genetic basis. In other words, <em>biological races do not exist in human populations. <\/em>However, the concept of race has been perpetuated and upheld by sociocultural constructs of race.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The conundrum for forensic anthropologists is the fact that while races do not exist on a biological level, we still socially recognize and categorize individuals based on their phenotype. Clearly, our phenotype is an important factor in not only how we are viewed by others but also how we identify ourselves. It is also a commonly reported variable. Often labeled as \u201crace,\u201d we are asked to report how we self-identify on school applications, government identification, surveys, census reports, and so forth. It follows then that when a person is reported missing, the information commonly collected by law enforcement and sometimes entered into a missing person\u2019s database includes their age, biological sex, stature, and \u201crace.\u201d Therefore, the more information a forensic anthropologist can provide regarding the individual\u2019s physical characteristics, the more he or she can help to narrow the search.<\/p>\n<p class=\"import-Normal\">As an exercise, create a list of all of the women you know who are between the ages of 18 and 24 and approximately 5\u2019 4\u201d to 5\u2019 9\u201d tall. You probably have several dozen people on the list. Now, consider how many females you know who are between the ages of 18 and 24, are approximately 5\u2019 4\u201d to 5\u2019 9\u201d tall, and are Vietnamese. Your list is going to be significantly shorter. That\u2019s how missing persons searches go as well. The more information you can provide regarding a decedent\u2019s phenotype, the fewer possible matches law enforcement are left to investigate. This is why population affinity has historically been included as a part of the biological profile.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Traditionally, population affinity was accomplished through a visual inspection of morphological variants of the skull (morphoscopics). These methods focused on elements of the facial skeleton, including the nose, eyes, and cheek bones. However, in an effort to reduce subjectivity, nonmetric cranial traits are now assessed within a statistical framework to help anthropologists better interpret their distribution among living populations (Hefner and Linde 2018). Based on the observable traits, a macromorphoscopic analysis will allow the practitioner to create a statistical prediction of geographic origin. In essence, forensic anthropologists are using human variation in the estimation of geographic origin, by referencing documented frequencies of nonmetric skeletal indicators or macromorphoscopic traits.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Population affinity is also assessed through metric analyses. The computer program Fordisc is an anthropological tool used to estimate different components of the biological profile, including ancestry, sex, and stature. When using Fordisc, skeletal measurements are input into the computer software, and the program employs multivariate statistical classification methods, including discriminant function analysis, to generate a statistical prediction for the geographic origin of unknown remains based on the comparison of the unknown to the reference samples in the software program. Fordisc also calculates the likelihood of the prediction being correct, as well as how typical the metric data is for the assigned group.<\/p>\n<h4 class=\"import-Normal\"><em>Estimating Age-at-Death<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Estimating age-at-death from the skeleton relies on the measurement of two basic physiological processes: (1) growth and development and (2) degeneration (or aging). From fetal development on, our bones and teeth grow and change at a predictable rate. This provides for relatively accurate age estimates. After our bones and teeth cease to grow and develop, they begin to undergo structural changes, or degeneration, associated with aging. This does not happen at such predictable rates and, therefore, results in less accurate or larger age-range estimations.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">During growth and development stages, two primary methods used for estimations of age of subadults (those under the age of 18) are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1240\">epiphyseal union<\/a><\/strong> and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1244\">dental development.<\/a><\/strong> Epiphyseal union<strong> (<\/strong>or <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1242\">epiphyseal fusion<\/a><\/strong>) refers to the appearance and closure of the epiphyseal plates between the primary centers of growth in a bone and the subsequent centers of growth (see Figure 15.7). Prior to complete union, the cartilaginous area between the primary and secondary centers of growth is also referred to as the growth plates (Schaefer, Black, and Scheuer 2009). Different areas of the skeleton have documented differences in the appearance and closure of epiphyses, making this a reliable method for aging subadult remains (SWGANTH 2013).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">As an example of its utility in the identification process, epiphyseal development was used to identify two subadult victims of a fatal fire in Flint, Michigan, in February 2010. The remains represented two young girls, ages three and four. Due to the intensity of the fire, the subadult victims were differentiated from each other through the appearance of the patella, the kneecap. The patella is a bone that develops within the tendon of the quadriceps muscle at the knee joint. The patella begins to form around three to four years of age (Cunningham, Scheuer, and Black 2016, 407\u2013409). In the example above, radiographs of the knees showed the presence of a patella in the four-year-old girl and the absence of a clearly discernible patella in the three-year-old.<\/p>\n<figure style=\"width: 358px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image18-2.png\" alt=\"Cranial cast of child with exposed maxilla and mandible to see developing dentition.\" width=\"358\" height=\"358\" \/><figcaption class=\"wp-caption-text\">Figure 15.12: Dental development in a subadult. Credit: <a href=\"https:\/\/boneclones.com\/product\/5-year-old-human-child-skull-with-mixed-dentition-exposed-BC-189\">5-year-old Human Child Skull with Mixed Dentition Exposed<\/a> by <a href=\"https:\/\/boneclones.com\/\">\u00a9BoneClones<\/a> is used by permission and available here is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Dental development begins during fetal stages of growth and continues until the complete formation and eruption of the adult third molars (if present). The first set of teeth to appear are called deciduous or baby teeth. Individuals develop a total of 20 deciduous teeth, including incisors, canines, and molars. These are generally replaced by adult dentition as an individual grows (Figure 15.12). A total of 32 teeth are represented in the adult dental arcade, including incisors, canines, premolars, and molars. When dental development is used for age estimations, researchers use both tooth-formation patterns and eruption schedules as determining evidence. For example, the crown of the tooth forms first followed by the formation of the tooth root. During development, an individual can exhibit a partially formed crown or a complete crown with a partially formed root. The teeth generally begin the eruption process once the crown of the tooth is complete. The developmental stages of dentition are one of the most reliable and consistent aging methods for subadults (Langley, Gooding, and Tersigni-Tarrant 2017, 176\u2013177).<\/p>\n<figure style=\"width: 403px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image22-3.png\" alt=\"Surfaces of three pubic symphyses: billowy (A) to more flat (B) to rough (C).\" width=\"403\" height=\"224\" \/><figcaption class=\"wp-caption-text\">Figure 15.13: Examples of degenerative changes to the pubic symphysis: (A) young adult; (B) middle adult; (C) old adult. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Example of the progression of degenerative changes to the pubic symphysis (Figure 15.14)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropo logy<\/a> by Ashley Kendell is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Original photos by Dr. Julie Fleischman used by permission. Pubic symphyses are curated in the Hartnett-Fulginiti donated skeletal collection. Donation and research consent was provided by next of kin.]<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Degenerative changes in the skeleton typically begin after 18 years of age, with more prominent changes developing after an individual reaches middle adulthood (commonly defined as after 35 years of age in osteology). These changes are most easily seen around joint surfaces of the pelvis, the cranial vault, and the ribs. In this chapter, we focus on the pubic symphysis surfaces of the pelvis and the sternal ends of the ribs, which show metamorphic changes from young adulthood to older adulthood. The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1262\">pubic symphysis<\/a> <\/strong>is a joint that unites the left and right halves of the pelvis. The surface of the pubic symphysis changes during adulthood, beginning as a surface with pronounced ridges (called billowing) and flattening with a more distinct rim to the pubic symphysis as an individual ages. As with all metamorphic age changes, older adults tend to develop lipping around the joint surfaces as well as a breakdown of the joint surfaces. The most commonly used method for aging adult skeletons from the pubic symphysis is the Suchey-Brooks method (Brooks and Suchey 1990; Katz and Suchey 1986). This method divides the changes seen with the pubic symphysis into six phases based on macroscopic age-related changes to the surface. Figure 15.13 provides a visual of the degenerative changes that typically occur on the pubic symphysis.<\/p>\n<figure style=\"width: 403px\" class=\"wp-caption alignright\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image12-3.png\" alt=\"Three sternal rib ends demonstrating progressive changes that occur with age.\" width=\"403\" height=\"220\" \/><figcaption class=\"wp-caption-text\">Figure 15.14: Examples of degenerative changes to the sternal rib end: (A) young adult; (B) middle adult; (C) old adult. Images derived from CSU, Chico HIL donated skeletal collection. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Examples of degenerative changes to the sternal rib end (Figure 15.15)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The sternal end of the ribs, the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1246\">anterior <\/a><\/strong> end of the rib that connects via cartilage to the sternum, is also used in age estimations of adults. This method, first developed by M. Y. \u0130\u015fcan and colleagues, considers both the change in shape of the sternal end as well as the quality of the bone (\u0130\u015fcan, Loth, and Wright 1984; \u0130\u015fcan, Loth, and Wright 1985). The sternal end first develops a billowing appearance in young adulthood. The bone typically develops a wider and deeper cupped end as an individual ages. Older adults tend to exhibit bony extensions of the sternal end rim as attaching cartilage ossifies. Figure 15.14 provides a visual of the degenerative changes that typically occur in sternal rib ends.<\/p>\n<h4 class=\"import-Normal\"><em>Estimating Stature<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Stature, or height, is one of the most prominently recorded components of the biological profile. Our height is recorded from infancy through adulthood. Doctor\u2019s appointments, driver's license applications, and sports rosters all typically involve a measure of stature for an individual. As such, it is also a component of the biological profile nearly every individual will have on record. Bioarchaeologists and forensic anthropologists use stature estimation methods to provide a range within which an individual\u2019s biological height would fall. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1248\">Biological height<\/a> <\/strong>is a person\u2019s true anatomical height. However, the range created through these estimations is often compared to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1264\">reported stature<\/a><\/strong>, which is typically self-reported and based on an approximation of an individual\u2019s true height (Ousley 1995).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In June 2015, two men were shot and killed in Granite Bay, California, in a double homicide. Investigators were able to locate surveillance camera footage from a gas station where the two victims were spotted in a car with another individual believed to be the perpetrator in the case. The suspect, sitting behind the victims in the car, hung his right arm out of the window as the car drove away. The search for the perpetrator was eventually narrowed down to two suspects. One suspect was 5\u2019 8\u201d while the other suspect was 6\u2019 4\u201d, representing almost a foot difference in height reported stature between the two. Forensic anthropologists were given the dimensions of the car (for proportionality of the arm) and were asked to calculate the stature of the suspect in the car from measurements of the suspect\u2019s forearm hanging from the window. Approximate lengths of the bones of the forearm were established from the video footage and used to create a predicted stature range. Stature estimations from skeletal remains typically look at the correlation between the measurements of any individual bone and the overall measurement of body height. In the case above, the length of the right forearm pointed to the taller of the two suspects who was subsequently arrested for the homicide.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Certain bones, such as the long bones of the leg, contribute more to our overall height than others and can be used with mathematical equations known as regression equations. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1252\">Regression methods <\/a> <\/strong>examine the relationship between variables such as height and bone length and use the correlation between the variables to create a prediction interval (or range) for estimated stature. This method for calculating stature is the most commonly used method (SWGANTH 2012). Figure 15.15 shows the measurement of the bicondylar length of the femur for stature estimations.<\/p>\n<figure style=\"width: 584px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image2-3.png\" alt=\"A femur is measured using a wooden osteometric board.\" width=\"584\" height=\"389\" \/><figcaption class=\"wp-caption-text\">Figure 15.15: Image of measurement of the bicondylar length of the femur, often used in the estimation of living stature. Image derived from CSU, Chico HIL donated skeletal collection. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Measurement of the bicondylar length of the femur<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Identification Using Individualizing Characteristics<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">One of the most frequently requested analyses within the forensic anthropology laboratory is assistance with the identification of unidentified remains. While all components of a biological profile, as discussed above, can assist law enforcement officers and medical examiners to narrow down the list of potential identifications, a biological profile will not lead to a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1254\">positive identification<\/a><\/strong>. The term <em>positive identification<\/em> refers to a scientifically validated method of identifying previously unidentified remains. Presumptive identifications, however, are not scientifically validated; rather, they are based on circumstances or scene context. For example, if a decedent is found in a locked home with no evidence of forced entry but the body is no longer visually identifiable, it may be presumed that the remains belong to the homeowner. Hence, a presumptive identification.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The medicolegal system ultimately requires that a positive identification be made in such circumstances, and a presumptive identification is often a good way to narrow down the pool of possibilities. Biological profile information also assists with making a presumptive identification based on an individual\u2019s phenotype in life (e.g., what they looked like). As an example, a forensic anthropologist may establish the following components of a biological profile: white male, between the ages of 35 and 50, approximately 5\u2019 7\u201d to 5\u2019 11.\u201d While this seems like a rather specific description of an individual, you can imagine that this description fits dozens, if not hundreds, of people in an urban area. Therefore, law enforcement can use the biological profile information to narrow their pool of possible identifications to include only white males who fit the age and height outlined above. Once a possible match is found, the decedent can be identified using a method of positive identification.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Positive identifications are based on what we refer to as individualizing traits or characteristics, which are traits that are unique at the individual level. For example, brown hair is not an individualizing trait as brown is the most common hair color in the U.S. But, a specific pattern of dental restorations or surgical implants can be individualizing, because it is unlikely that you will have an exact match on either of these traits when comparing two individuals.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">A number of positive methods are available to forensic anthropologists, and for the remainder of this section we will discuss the following methods: comparative medical and dental radiography and identification of surgical implants.<\/p>\n<figure style=\"width: 165px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image17-3.png\" alt=\"Radiograph of skull with frontal sinuses visible.\" width=\"165\" height=\"182\" \/><figcaption class=\"wp-caption-text\">Figure 15.16: Example of the unique shape of the frontal sinus. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Frontal_bone_sinuses.jpg\">Frontal bone sinuses<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Alex_Khimich\">Alex Khimich<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Comparative medical and dental radiography is used to find consistency of traits when comparing antemortem records (medical and dental records taken during life) with images taken postmortem (after death). Comparative medical radiography focuses primarily on features associated with the skeletal system, including trabecular pattern (internal structure of bone that is honeycomb in appearance), bone shape or cortical density (compact outer layer of bone), and evidence of past trauma, skeletal pathology, or skeletal anomalies. Other individualizing traits include the shape of various bones or their features, such as the frontal sinuses (Figure 15.16).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Comparative dental radiography focuses on the number, shape, location, and orientation of dentition and dental restorations in antemortem and postmortem images. While there is not a minimum number of matching traits that need to be identified for an identification to be made, the antemortem and postmortem records should have enough skeletal or dental consistencies to conclude that the records did in fact come from the same individual (SWGANTH 2010a). Consideration should also be given to population-level frequencies of specific skeletal and dental traits. If a trait is particularly common within a given population, it may not be a good trait to utilize for positive identification.<\/p>\n<figure style=\"width: 354px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image16-3.png\" alt=\"A scapula and humerus with a metal shoulder replacement.\" width=\"354\" height=\"231\" \/><figcaption class=\"wp-caption-text\">Figure 15.17: Image of joint replacement in the right shoulder. Credit: <a href=\"https:\/\/naturalhistory.si.edu\/education\/teaching-resources\/written-bone\/skeleton-keys\/todays-bones\">Shoulder replacement<\/a> by <a href=\"https:\/\/www.si.edu\/\">Smithsonian<\/a> [exhibit: Written in Bone, Today\u2019s Bones] <a href=\"https:\/\/www.si.edu\/termsofuse\">is used for educational and non-commercial purposes as outlined by the Smithsonian.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Surgical implants or devices can also be used for identification purposes (Figure 15.17). These implements are sometimes recovered with human remains. One of the ways forensic anthropologists can use surgical implants to assist in decedent identification is by providing a thorough analysis of the implant and noting any identifying information such as serial numbers, manufacturer symbols, and so forth. This information can then sometimes be tracked directly to the manufacturer or the place of surgical intervention, which may be used to identify unknown remains (SWGANTH 2010a).<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Trans Doe Task Force<\/h2>\n<p class=\"import-Normal\">The Trans Doe Task Force (TDTF) is a Trans-led nonprofit organization that investigates cases involving LGBTQ+ missing and murdered persons. The organization specifically focuses on transgender and gender-variant cases, providing connections between law enforcement agencies, medical examiner offices, forensic anthropologists, and forensic genetic genealogists to increase the chances of identification. Additionally, the TDTF curates a data repository of missing, murdered, and unclaimed LGBTQ+ individuals, and they continuously try innovative approaches to identify these individuals, whose lived gender identity may not match their biological sex.<\/p>\n<p class=\"import-Normal\">For more information visit <a href=\"https:\/\/transdoetaskforce.org\/\">transdoetaskforce.org<\/a><\/p>\n<\/div>\n<h3 class=\"import-Normal\"><strong>Trauma Analysis<\/strong><\/h3>\n<h4 class=\"import-Normal\"><em>Types of Trauma<\/em><strong><br \/>\n<\/strong><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Within the field of anthropology, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1256\">trauma<\/a> <\/strong>is defined as an injury to living tissue caused by an extrinsic force or mechanism (Lovell 1997:139). Forensic anthropologists can assist a forensic pathologist by providing an interpretation of the course of events that led to skeletal trauma. Typically, traumatic injury to bone is classified into one of four categories, defined by the trauma mechanism. A trauma mechanism refers to the force that produced the skeletal modification and can be classified as (1) sharp force, (2) blunt force, (3) projectile, or (4) thermal (burning). Each type of trauma, and the characteristic pattern(s) associated with that particular categorization, will be discussed below.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">First, let\u2019s consider s<em>harp-force trauma<\/em>, which is caused by a tool that is edged, pointed, or beveled\u2014for example, a knife, saw, or machete (SWGANTH 2011). The patterns of injury resulting from sharp-force trauma include linear incisions created by a sharp, straight edge; punctures; and chop marks (Figure 15.18; SWGANTH 2011). When observed under a microscope, an anthropologist can often determine what kind of tool created the bone trauma. For example, a power saw cut will be discernible from a manual saw cut.<\/p>\n<figure style=\"width: 602px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image21-1.png\" alt=\"Anterior image of a skull with multiple traumatic injuries to forehead.\" width=\"602\" height=\"457\" \/><figcaption class=\"wp-caption-text\">Figure 15.18: Example of sharp-force trauma (sword wound) to the frontal bone. The skull appears sliced with thin lines in two places across the top of the skull. Credit: <a href=\"https:\/\/openverse.org\/image\/909d1b77-ad5f-4cda-be44-6d9b5fbf14b9\/\">Female skull injured by a medieval sword<\/a> by <a href=\"https:\/\/sketchfab.com\/provinciaal_depot_noordholland\">Provinciaal depot voor archeologie Noord-Holland<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY 4.0 License<\/a>. The original image is a 3D model that can be manipulated on the <a href=\"https:\/\/wordpress.org\/openverse\/image\/909d1b77-ad5f-4cda-be44-6d9b5fbf14b9\/\">openverse website<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Second, <em>blunt-force trauma<\/em> is defined as \u201ca relatively low-velocity impact over a relatively large surface area\u201d (Galloway 1999, 5). Blunt-force injuries can result from impacts from clubs, sticks, fists, and so forth. Blunt-force impacts typically leave an injury at the point of impact but can also lead to bending and deformation in other regions of the bone. Depressions, fractures, and deformation at and around the site of impact are all characteristics of blunt-force trauma (Figure 15.19). As with sharp-force trauma, an anthropologist attempts to interpret blunt-force injuries, providing information pertaining to the type of tool used, the direction of impact, the sequence of impacts, if more than one, and the amount of force applied.<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 578px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image30.png\" alt=\"Cranium with two blunt force impacts from a hammer.\" width=\"578\" height=\"803\" \/><figcaption class=\"wp-caption-text\">Figure 15.19: Example of multiple blunt force impacts to the left parietal and frontal bones. There is one hole in the skull with fractured bone around the edges. There are also multiple spots across the back of the skull with depressions of various sizes. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Skull_hammer_trauma.jpg\">Skull hammer trauma<\/a> by <a href=\"https:\/\/www.nih.gov\/\">the National Institutes of Health<\/a>, Health &amp; Human Services, is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>. [Exhibit: Visible Proofs: Forensic Views of the Body, U.S. National Library of Medicine, 19th Century Collection, National Museum of Health and Medicine, Armed Forces Institute of Pathology, Washington, D.C.]<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Third, <em>projectile trauma<\/em> refers to high-velocity trauma, typically affecting a small surface area (Galloway 1999, 6). Projectile trauma results from fast-moving objects such as bullets or shrapnel. It is typically characterized by penetrating defects or embedded materials (Figure 15.20). When interpreting injuries resulting from projectile trauma, an anthropologist can often offer information pertaining to the type of weapon used (e.g., rifle vs. handgun), relative size of the bullet (but not the caliber of the bullet), the direction the projectile was traveling, and the sequence of injuries if there are multiple present.<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 462px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image5-3.png\" alt=\"Anterior and posterior views of a skull with a gunshot wound.\" width=\"462\" height=\"291\" \/><figcaption class=\"wp-caption-text\">Figure 15.20: Example of projectile trauma with an entrance wound to the frontal bone and exit wound visible on the occipital. A small circular hole is visible in the front of the skull with cracks radiating out from the point of impact. There is a larger hole visible in the back of the skull that is irregular yet circular in shape. Credit: <a href=\"https:\/\/naturalhistory.si.edu\/education\/teaching-resources\/written-bone\/skeleton-keys\/how-bone-biographies-get-written\">Trauma: Gunshot Wounds<\/a> by <a href=\"https:\/\/www.si.edu\/\">Smithsonian<\/a> [exhibit: Written in Bone, How Bone Biographies Get Written] <a href=\"https:\/\/www.si.edu\/termsofuse\">is used for educational and non-commercial purposes as outlined by the Smithsonian.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Finally, <em>thermal trauma<\/em> is a bone alteration that results from bone exposure to extreme heat. Thermal trauma can result in cases of house or car fires, intentional disposal of a body in cases of homicidal violence, plane crashes, and so on. Thermal trauma is most often characterized by color changes to bone, ranging from yellow to black (charred) or white (calcined). Other bone alterations characteristic of thermal trauma include delamination (flaking or layering due to bone failure), shrinkage, fractures, and heat-specific burn patterning. When interpreting injuries resulting from thermal damage, an anthropologist can differentiate between thermal fractures and fractures that occurred before heat exposure, thereby contributing to the interpretation of burn patterning (e.g., was the individual bound or in a flexed position prior to the fire?).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">While there are characteristic patterns associated with the four categories of bone trauma, it is also important to note that these bone alterations do not always occur independently of different trauma types. An individual\u2019s skeleton may present with multiple different types of trauma, such as a projectile wound and thermal trauma. Therefore, it is important that the anthropologist recognize the different types of trauma and interpret them appropriately.<\/p>\n<h3 class=\"import-Normal\"><strong>Timing of Injury<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Another important component of any anthropological trauma analysis is the determination of the timing of injury (e.g., when did the injury occur). Timing of injury is traditionally split into one of three categories: <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1260\">antemortem<\/a> <\/strong>(before death), <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1258\">perimortem<\/a> <\/strong>(at or around the time of death), and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1266\">postmortem <\/a><\/strong>(after death). This classification system differs slightly from the classification system used by the pathologist because it specifically references the qualities of bone tissue and bone response to external forces. Therefore, the perimortem interval (at or around the time of death) means that the bone is still fresh and has what is referred to as a green bone response, which can extend past death by several weeks or even months. For example, in cold or freezing temperatures a body can be preserved for extended periods of time, increasing the perimortem interval, while in desert climates decomposition is accelerated, thereby significantly decreasing the postmortem interval (Galloway 1999, 12). Antemortem injuries (occurring well before death and not related to the death incident) are typically characterized by some level of healing, in the form of a fracture callus or unification of fracture margins. Finally, postmortem injuries (occurring after death, while bone is no longer fresh) are characterized by jagged fracture margins, resulting from a loss of moisture content during the decomposition process (Galloway 1999, 16). In general, all bone traumas should be classified according to the timing of injury, if possible. This information will help the medical examiner or pathologist better understand the circumstances surrounding the decedent\u2019s death, as well as events occurring during life and after the final disposition of the body.<\/p>\n<h3 class=\"import-Normal\"><strong>The Role of the Forensic Anthropologist in Trauma Analysis<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Within the medicolegal system, forensic anthropologists are often called upon by the medical examiner, forensic pathologist, or coroner to assist with an interpretation of trauma. The forensic anthropologist\u2019s main focus in any trauma analysis is the underlying skeletal system\u2014as well as, sometimes, cartilage. Analysis and interpretation of soft tissue injuries fall within the purview of the medical examiner or pathologist. It is also important to note that the main role of the forensic anthropologist is to provide information pertaining to skeletal injury to assist the medical examiner\/pathologist in their final interpretation of injury. Forensic anthropologists do not hypothesize as to the cause of death of an individual. Instead, a forensic anthropologist\u2019s report should include a description of the injury (e.g., trauma mechanism, number of injuries, location, timing of injury); documentation of the injury, which may be utilized in court testimony (e.g., photographs, radiographs, measurements); and, if applicable, a statement as to the condition of the body and state of decomposition, which may be useful for understanding the depositional context (e.g., how long has the body been exposed to the elements; was it moved or in its original location; are any of the alterations to bone due to environmental or faunal exposure instead of intentional human modification).<\/p>\n<h2 class=\"import-Normal\">Taphonomy<\/h2>\n<h2 class=\"import-Normal\"><strong>What Happened to the Remains After Death?<\/strong><\/h2>\n<p class=\"import-Normal\">The majority of the skeletal analysis process revolves around the identity of the deceased individual. However, there is one last, very important question that forensic anthropologists should ask: What happened to the remains after death? Generally speaking, processes that alter the bone after death are referred to as taphonomic changes (refer to Chapter 7 for a discussion regarding taphonomy and the fossil record).<\/p>\n<p class=\"import-Normal\">The term <em>taphonomy<\/em> was originally used to refer to the processes through which organic remains mineralize, also known as fossilization. Within the context of biological anthropology, the term <em>taphonomy<\/em> is better defined as the study of what happens to human remains after death (Komar and Buikstra 2008). Initial factors affecting a body after death include processes such as decomposition and scavenging by animals. However, taphonomic processes encompass much more than the initial period after death. For example, plant root growth can leach minerals from bone, leaving a distinctive mark. Sunlight can bleach human remains, leaving exposed areas whiter than those that remained buried. Water can wear the surface of the bone until it becomes smooth.<\/p>\n<p class=\"import-Normal\">Some taphonomic processes can help a forensic anthropologist estimate the relative amount of time that human remains have been exposed to the elements. For example, root growth through a bone would certainly indicate a body was buried for more than a few days. Forensic anthropologists must be very careful when attempting to estimate time since death based on taphonomic processes because environmental conditions can greatly influence the rate at which taphonomic processes progress. For example, in cold environments, tissue may decay slower than in warm, moist environments.<\/p>\n<p class=\"import-Normal\">Forensic anthropologists must contend with taphonomic processes that affect the preservation of bones. For example, high acidity in the soil can break down human bone to the point of crumbling. In addition, when noting trauma, they must be very careful not to confuse postmortem (after death) bone damage with trauma.<\/p>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 470.25pt\">\n<caption>Figure 15.21: Table showing taphonomic processes that affect the preservation of bones. A. Rodent gnawing. B. Carnivore damage. C. Burned bone. D. Root etching. E. Weathering. F. Cut marks. Credit: A. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Rodent gnawing (Figure 15.26)<\/a>, B. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Carnivore damage (Figure 15.27)<\/a>, C. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Burned bone (Figure 15.28)<\/a>, D. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Root etching (Figure 15.29)<\/a>, E. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Weathering (Figure 15.30)<\/a>, and F. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-5\/\">Cut marks (Figure 15.30)<\/a>, all original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Alex Perrone are under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-NC 4.0 License<\/a>.<\/caption>\n<thead>\n<tr style=\"height: 52.5pt\">\n<td class=\"Table1-C\" style=\"padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Taphonomic Process<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 1pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Definition<\/p>\n<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table1-R\" style=\"height: 190.5pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Rodent Gnawing<\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: center\"><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image19-2.png\" alt=\"Parallel tooth marks etched by a rodent\u2019s front teeth visible on the end of an animal bone.\" width=\"564\" height=\"422\" \/><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">When rodents, such as rats and mice, chew on bone, they leave sets of parallel grooves. The shallow grooves are etched by the rodent\u2019s incisors.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 166.75pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Carnivore Damage<\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: center\"><strong><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image23-4.png\" alt=\"Pit marks from the canines of a carnivore visible on the surface of an animal bone.\" width=\"410\" height=\"272\" \/><\/strong><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Carnivores may leave destructive dental marks on bone. The tooth marks may be visible as pit marks or punctures from the canines, as well as extensive gnawing or chewing of the ends of the bones to retrieve marrow.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 177pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Burned Bone<\/strong><\/p>\n<p class=\"import-Normal\"><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image20-5.png\" alt=\"Burned animal bone fragments pictured at different stages of thermal damage.\" width=\"512\" height=\"342\" \/><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Fire causes observable damage to bone. Temperature and the amount of time bone is heated affect the appearance of the bone. Very high temperatures can crack bone and result in white coloration. Color gradients are visible in between high and lower temperatures, with lower temperatures resulting in black coloration from charring. Cracking can also reveal information about the directionality of the burn.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 169.75pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Root Etching<\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: center\"><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image15-4.png\" alt=\"Animal bone with prominent, discolored grooves where roots leached nutrients from bone\u2019s surface.\" width=\"512\" height=\"342\" \/><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Plant roots can etch the outer surface of bone, leaving grooves where the roots attached as they leached nutrients. During this process, the plant\u2019s roots secrete acid that breaks down the surface of the bone.<\/p>\n<p class=\"import-Normal\">\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 170.5pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Weathering<\/strong><\/p>\n<p class=\"import-Normal\"><strong><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image9.png\" alt=\"Cracking and exfoliation of the surface of an animal bone. \" width=\"512\" height=\"342\" \/><\/strong><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Many different environmental conditions affect bone. River transport can smooth the surface of the bone due to water abrasion. Sunlight can bleach the exposed surface of bone. Dry and wet environments or the mixture of both types of environments can cause cracking and exfoliation of the surface. Burial in different types of soil can cause discoloration, and exposure can cause degreasing.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 169.75pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 1pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\" style=\"text-align: center;margin-left: 36pt\"><strong>Cut Marks<\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: left\"><img class=\"alignnone\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image8-2.png\" alt=\"Thin vertical lines and cuts are visible along the bone.\" width=\"512\" height=\"342\" \/><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.75pt;border-right: solid #000000 1pt;border-bottom: solid #000000 1pt;border-left: solid #000000 0.75pt;padding: 5pt 5pt 5pt 5pt\">\n<p class=\"import-Normal\">Humans may alter bone by cutting, scraping, or sawing it directly or in the process of removing tissue. The groove pattern\u2014that is, the depth and width of the cuts\u2014can help identify the tool used in the cutting process.<\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div class=\"textbox kk shaded\">\n<h2>Dig Deeper: Modern Forensic Technologies<\/h2>\n<p>In recent years, the forensics community has greatly benefited from the introduction of new technologies, helping strengthen the precision and speed of discoveries and advancements in the field. With recent developments in forensic anthropology, such as 3D scanning technologies, virtual reconstruction, and AI-assisted DNA analysis being integrated into traditional methods, there have been notable changes in how experts investigate human remains.<\/p>\n<p><strong>Artificial intelligence<\/strong><\/p>\n<p>In recent years, Artificial intelligence (AI) has shown itself to be a valuable tool within forensic anthropology. Aiding forensic experts and toxicologists with complex tasks, the limitations of traditional autopsies can be addressed with the help of AI. By automating and enhancing key investigative processes such as searching for microscopic changes in the human body to determine the cause of death or a person\u2019s life conditions, AI has the potential to enhance the efficiency of forensic processes significantly. It facilitates the detection of microscopic bodily changes to determine the cause of death or living conditions, compares evidence against databases for weapon identification and blood spatter analysis, and reduces manual workload. AI also enables the electronic storage of biometric data\u2013such as facial features, retinal patterns, and fingerprints\u2013for more accurate identity verification. Additionally, AI-powered microscopy enhances the detection of biological traces on complex surfaces, while blood biomarker analysis allows for more precise estimations of time of death (Wankhade et al., 2022).<\/p>\n<p>While AI holds great promise for the future of forensic medicine, a significant challenge remains: sourcing high-quality data to train the algorithms effectively. One of the more recent AI technologies making waves in the forensic anthropology sector is a new automated AI algorithm called the Convolutional Neural Network (CNN). As described by researchers in Switzerland\u2019s national medical journal Healthcare, CNN is a Deep Learning algorithm that allows for the detection of microscopic skull damage from CT scans or soft-tissue predictions of a face based on the skull information provided (Thurzo et al., 2021). While there are many advantages to using the CNN, the algorithm can be subject to biases in the same way human forensic experts can, as its assessment and pattern recognition of skulls and skeletons depend on the source data initially used for its AI training (2021).<\/p>\n<p><strong>3D Modeling<\/strong><\/p>\n<p>Identifying complex trauma to bones\u2013such as distinguishing heat fractures following blunt force trauma\u2013remains a significant challenge in forensic anthropology. This is particularly true for irregular skeletal structures like the pelvis, where overlapping trauma types can be difficult to differentiate, leading to these bones often being understudied. A 2024 study done by researchers from the University of Alberta in collaboration with the Michigan State Police explores the use of 3D laser scans and modelling technology to provide a highly detailed analysis of irregular bones with trauma. The study aimed to better distinguish peri-mortem trauma (trauma occurring around the time of death) from post-mortem heat alterations and improve the forensic analysis accuracy of such cases (Friedlander et al., 2024). The use of 3D laser scans and modelling technology provides very clear, detailed, and colored scans of bones, showing distinctions between the characteristics of the fractures. Blunt force and sharp force trauma produce a colour gradient on the 3D model that is more gradual and irregular, while heat fractures are more neat and characterized by little colour variation on the 3D models (2024). Other conclusions were also drawn from the study, such as the differences in trauma on fresh bones and bones that have been exposed to the elements for longer. An example of this is the interstitial fluid and collagen fibrils in fresh bones absorbing force, causing more long and jagged fracture lines, as opposed to a brittle fracture that older bones may exhibit (2024).<\/p>\n<p>Overall, the integration of 3D modeling technology offers a reproducible and highly detailed approach for analyzing trauma in anatomically complex and historically understudied skeletal regions. The practicality of this advancement is further emphasized by the researchers, who note that \u201cin many instances, scanned 3D models can be 3D printed for handheld representation of the model without damaging or overhandling the remains\u201d (2024, p. 2). By enhancing the ability to differentiate between various types of trauma and allowing for more convenient and risk-averse methods of research, this technology significantly improves the accuracy and reliability of forensic interpretations.<\/p>\n<\/div>\n<h2>Ethics and Human Rights<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Working with human remains requires a great deal of consideration and respect for the dead. Forensic anthropologists have to think about the ethics of our use of human remains for scientific purposes. How do we conduct casework in the most respectable manner possible? While there are a wide range of ethical considerations to consider when contemplating a career in forensic anthropology, this chapter will focus on two major categories: working with human remains and acting as an expert within the medicolegal system.<\/p>\n<h3 class=\"import-Normal\"><strong>Working with Human Remains<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropologists work with human remains in a number of contexts, including casework, excavation, research, and teaching. When working with human remains, it is always important to use proper handling techniques. To prevent damage to skeletal remains, bones should be handled over padded surfaces. Skulls should never be picked up by placing fingers in the eye orbits, foramen magnum (hole at the base of the skull for entry of the spinal cord), or through the zygomatic arches (cheekbones). Human remains, whether related to casework, fieldwork, donated skeletal collections, or research, were once living human beings. It is important to always bear in mind that work with remains should be ingrained with respect for the individual and their relatives. In addition to fieldwork, casework, and teaching, anthropologists are often invited to work with remains that come from a bioarchaeological context or from a human rights violation. While this discussion of ethics is not comprehensive, two case examples will be provided below in which an anthropologist must consider the ethical standards outlined above.<\/p>\n<h3 class=\"import-Normal\"><strong>Modern Human Rights Violations<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Forensic anthropologists may also be called to participate in criminal investigations involving human rights violations. Anthropological investigations may include assistance with identifications, determination of the number of victims, and trauma analyses. In this role, forensic anthropologists play an integral part in promoting human rights, preventing future human rights violations, and providing the evidence necessary to prosecute those responsible for past events. A few ethical considerations for the forensic anthropologist involved in human rights violations include the use of appropriate standards of identification, presenting reliable and unbiased testimony, and maintaining preservation of evidence. For a more comprehensive history of forensic anthropological contributions to human rights violations investigations, see Ubelaker 2018.<\/p>\n<h3 class=\"import-Normal\"><strong>Acting as an Expert in the Medicolegal System<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In addition to the ethical considerations involved in working with human skeletal remains, forensic anthropologists must abide by ethical standards when they act as experts within the medicolegal system. The role of the forensic anthropologist within the medicolegal system is primarily to provide information to the medical examiner or coroner that will aid in the identification process or determination of cause and manner of death. Forensic anthropologists also may be called to testify in a court of law. In this capacity, forensic anthropologists should always abide by a series of ethical guidelines that pertain to their interpretation, presentation, and preservation of evidence used in criminal investigations. First and foremost, practitioners should never misrepresent their training or education. When appropriate, outside opinions and assistance in casework should be requested (e.g., consulting a radiologist for radiological examinations or odontologist for dental exams). The best interest of the decedent should always take precedence. All casework should be conducted in an unbiased way, and financial compensation should never be accepted as it can act as an incentive to take a biased stance regarding casework. All anthropological findings should be kept confidential, and release of information is best done by the medical examiner or coroner. Finally, while upholding personal ethical standards, forensic anthropologists are also expected to report any perceived ethical violations committed by their peers.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ethical standards for the field of forensic anthropology are outlined by the Organization of Scientific Area Committees (OSAC) for Forensic Science, administered by the National Institute of Standards and Technology (NIST). OSAC and NIST recently began an initiative to develop standards that would strengthen the practice of forensic science both in the United States and internationally. OSAC\u2019s main objective is to \u201cstrengthen the nation\u2019s use of forensic science by facilitating the development of technically sound forensic science standards and by promoting the adoption of those standards by the forensic science community\u201d (NIST n.d.). Additionally, OSAC promotes the establishment of best practices and other guidelines to ensure that forensic science findings and their presentation are reliable and reproducible (NIST 2023).<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Native American Graves Protection and Repatriation Act (NAGPRA)<\/h2>\n<p class=\"import-Normal\">There is a long history in the <span style=\"background-color: #00ffff\">United States<\/span> of systematic disenfranchisement of Native American people, including lack of respect for tribal sovereignty. This includes the egregious treatment of Native American human remains. Over several centuries, thousands of Native American remains were removed from tribal lands and held at institutions in the United States, such as museums and universities.<\/p>\n<p class=\"import-Normal\">In 1990, a landmark human rights federal law, the Native American Graves Protection and Repatriation Act (NAGPRA), spurred change in the professional standards and practice of biological anthropology and archaeology. NAGPRA established a legal avenue to provide protection for and repatriation of Native American remains, cultural items, and sacred objects removed from Federal or tribal lands to Native American lineal descendants, Indian tribes, and Native Hawaiian organizations. Human remains and associated artifacts, curated in museum collections and federally funded institutions, are subject to three primary provisions outlined by the NAGPRA statute: (1) protection for Native graves on federal and private land; (2) recognition of tribal authority on such lands; and (3) the requirement that all Native skeletal remains and associated artifacts be inventoried and culturally affiliated groups be consulted concerning decisions related to ownership and final disposition (Rose, Green, and Green 1996). NAGPRA legislation was enacted to ensure ethical consideration and treatment of Native remains and to improve dialogue between scientists and Native groups.<\/p>\n<ul>\n<li>For more information about NAGPRA, visit the <a href=\"https:\/\/www.usbr.gov\/nagpra\/\" target=\"_blank\" rel=\"noopener\">Bureau of Reclamation NAGPRA website<\/a><\/li>\n<li>To read the text of the law, visit the <a href=\"https:\/\/www.congress.gov\/bill\/101st-congress\/house-bill\/5237\">US Congress NAGPRA law website<\/a>.<\/li>\n<li>For further discussion of NAGPRA history, please see <a href=\"https:\/\/textbooks.whatcom.edu\/tracesarchaeology\/\" target=\"_blank\" rel=\"noopener\"><em>TRACES: <\/em><em>An Open Invitation to <\/em><em>Archaeology <\/em>open textbook website<\/a><em><br \/>\n<\/em><\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\"><span style=\"background-color: #ff99cc\">Becoming a Forensic Anthropologist<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"background-color: #ff99cc\">What does it take to be a forensic anthropologist? Forensic anthropologists are first and foremost anthropologists. While many forensic anthropologists have an undergraduate degree in anthropology, they may also major in biology, criminal justice, pre-law, pre-med, and many other related fields. Practicing forensic anthropologists typically have an advanced degree, either a Master\u2019s or Doctoral degree in Anthropology. Additional training and experience in archaeology, the medico-legal system, rules of evidence, and expert witness testimony are also common. Practicing forensic anthropologists are also encouraged to be board-certified through the American Board of Forensic Anthropology (ABFA). Learn more about the field and educational opportunities on the ABFA website: <a class=\"rId111\" style=\"background-color: #ff99cc\" href=\"https:\/\/www.theabfa.org\/coursework\">https:\/\/www.theabfa.org\/coursework<\/a>.<\/span><\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Review Questions<\/h2>\n<ul>\n<li>What is forensic anthropology? What are the seven primary steps involved in a skeletal analysis?<\/li>\n<li>What are the major components of a biological profile? Why are forensic anthropologists often-tasked with creating biological profiles for unknown individuals?<\/li>\n<li>What are the four major types of skeletal trauma?<\/li>\n<li>What is taphonomy, and why is an understanding of taphonomy often critical in forensic anthropology analyses?<\/li>\n<li>What are some of the ethical considerations faced by forensic anthropologists?<\/li>\n<\/ul>\n<\/div>\n<h2>About the Authors<\/h2>\n<p><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image24-4.jpg\" alt=\"A woman with straight blonde hair smiles at the camera. \" width=\"191\" height=\"254\" \/><\/p>\n<h3 class=\"import-Normal\"><strong>Ashley Kendell, Ph.D.<\/strong><\/h3>\n<p class=\"import-Normal\">California State University, Chico, <a class=\"rId113\" href=\"mailto:akendell@csuchico.edu\">akendell@csuchico.edu<\/a><\/p>\n<p class=\"import-Normal\">Dr. Ashley Kendell is currently an associate professor and forensic anthropologist at Chico State. Prior to beginning her position at Chico State, she was a visiting professor at the University of Montana and the forensic anthropologist for the state of Montana. Dr. Kendell obtained her doctorate from Michigan State University, and her research interests include skeletal trauma analysis and digitization and curation methods for digital osteological data. She is also a Registry Diplomate of the American Board of Medicolegal Death Investigators. Throughout her doctoral program, she worked as a medicolegal death investigator for the greater Lansing, Michigan, area and was involved in the investigation of over 200 forensic cases.<\/p>\n<p class=\"import-Normal\"><strong><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image4.jpg\" alt=\"A woman with straight brown hair pulled back smiles at the camera. \" width=\"194\" height=\"258\" \/><\/strong><\/p>\n<h3 class=\"import-Normal\"><strong>Alex Perrone, M.A., M.S.N, R.N., P.H.N.<\/strong><\/h3>\n<p class=\"import-Normal\">Butte Community College, <a class=\"rId115\" href=\"mailto:perroneal@butte.edu\">perroneal@butte.edu<\/a><\/p>\n<p class=\"import-Normal\">Alex Perrone is a lecturer in anthropology at Butte Community College. She is also a Registered Nurse and a certified Public Health Nurse. She is a former Supervisor of the Human Identification Laboratory in the Department of Anthropology at California State University, Chico. Her research interests include bioarchaeology, paleopathology, forensic anthropology, skeletal biology, California prehistory, and public health. She has worked on bioarchaeological and archaeological projects in Antigua, California, Hawaii, Greece, and the UK, and was an archaeological technician for the USDA Forest Service. She assisted with training courses for local and federal law enforcement agencies and assisted law enforcement agencies with the recovery and analysis of human remains.<\/p>\n<p class=\"import-Normal\" data-wp-editing=\"1\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image1-1.jpg\" alt=\"A woman with curly brown, shoulder-length hair smiles at the camera.\" width=\"190\" height=\"253\" \/><\/p>\n<h3 class=\"import-Normal\"><strong>Colleen Milligan, Ph.D.<\/strong><\/h3>\n<p class=\"import-Normal\">California State University, Chico, <a class=\"rId117\" href=\"mailto:cfmilligan@csuchico.edu\">cfmilligan@csuchico.edu<\/a><\/p>\n<p class=\"import-Normal\">Dr. Colleen Milligan is a biological and forensic anthropologist with research interests in bioarchaeology, skeletal biology, and forensic anthropology. She has been a Fellow with the Department of Homeland Security and has assisted in forensic anthropology casework and recoveries in the State of Michigan and California. She has also assisted in community outreach programs in forensic anthropology and forensic science, as well as recovery training courses for local, state, and federal law enforcement officers. She is a certified instructor through Peace Officers Standards and Training (POST). Dr. Milligan serves as the current co-director of the Chico State Human Identification Laboratory.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration<\/h2>\n<p><a href=\"https:\/\/www.theabfa.org\/coursework\" target=\"_blank\" rel=\"noopener\">The American Board of Forensic Anthropology (ABFA)<\/a><\/p>\n<p><a href=\"https:\/\/www.aafs.org\/\" target=\"_blank\" rel=\"noopener\">The American Academy of Forensic Sciences (AAFS)<\/a><\/p>\n<p><a href=\"https:\/\/www.nist.gov\/organization-scientific-area-committees-forensic-science\" target=\"_blank\" rel=\"noopener\">The Organization of Scientific Area Committees for Forensic Science (OSAC)<\/a><\/p>\n<p><a href=\"https:\/\/textbooks.whatcom.edu\/tracesarchaeology\/\" target=\"_blank\" rel=\"noopener\">TRACES Bioarchaeology<\/a><\/p>\n<p><a href=\"https:\/\/transdoetaskforce.org\/\" target=\"_blank\" rel=\"noopener\">Trans Doe Task Force<\/a><\/p>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Adams, Bradley J., and Lyle W. Konigsberg, eds. 2008. <em>Recovery, Analysis, and Identification of Commingled Remains<\/em>. Totowa, NJ: Humana Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Beatrice, Jared S., and Angela Soler. 2016. \u201cSkeletal Indicators of Stress: A Component of the Biocultural Profile of Undocumented Migrants in Southern Arizona.\u201d <em>Journal of Forensic Sciences <\/em>61 (5): 1164\u20131172.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Berg, Gregory E. 2017. \u201cSex Estimation of Unknown Human Skeletal Remains.\u201d In <em>Forensic Anthropology: A Comprehensive Introduction, Second Edition<\/em>, edited by Natalie R. Langley and MariaTeresa A. Tersigni-Tarrant, 143\u2013159. Boca Raton, FL: CRC Press.<\/p>\n<p class=\"import-Normal\">Bethard, Jonathan D., and Elizabeth A. DiGangi. 2020. \u201cLetter to the Editor\u2014Moving Beyond a Lost Cause: Forensic Anthropology and Ancestry Estimates in the United States.\u201d <em>Journal of Forensic Sciences<\/em> 65 (5): 1791\u20131792.<\/p>\n<p class=\"import-Normal\">Birkby, Walter H., Todd W. Fenton, and Bruce E. Anderson. 2008. \u201cIdentifying Southwest Hispanics Using Nonmetric Traits and the Cultural Profile.\u201d <em>Journal of Forensic Sciences <\/em>53 (1): 29\u201333.<\/p>\n<p class=\"import-Normal\">Blatt, Samantha, Amy Michael, and Lisa Bright. Forthcoming. \u201cBioarchaeology: Interpreting Human Behavior from Skeletal Remains.\u201d In <em>TRACES: <\/em><em>An Open Invitation to <\/em><em>Archaeology<\/em>. https:\/\/textbooks.whatcom.edu\/tracesarchaeology\/.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Brooks, S., and J. M. Suchey. 1990. \u201cSkeletal Age Determination Based on the Os Pubis: A Comparison of the Acs\u00e1di-Nemesk\u00e9ri and Suchey-Brooks Methods.\u201d <em>Human Evolution <\/em>5 (3): 227\u2013238.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Buchanan, Shelby. 2014. \u201cBone Modification in Male to Female Transgender Surgeries: Considerations for the Forensic Anthropologist.\u201d MA thesis, Department of Geography and Anthropology, Louisiana State University, Baton Rouge.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Cunningham, Craig, Louise Scheuer, and Sue Black. 2016. <em>Developmental Juvenile Osteology, Second Edition<\/em>. London: Elsevier Academic Press.<\/p>\n<p>Friedlander, H., Adeeb, S., Correia, P. M., Stone, D., &amp; Brooks\u2010Lim, E. (2024). An innovative way to use 3d modeling on burnt bone to differentiate heat fractures from blunt and sharp force trauma. WIREs Forensic Science, 6(5), 1\u201318. <a href=\"https:\/\/doi.org\/10.1002\/wfs2.1525\">https:\/\/doi.org\/10.1002\/wfs2.1525<\/a><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Galloway, Alison, ed. 1999. <em>Broken Bones: Anthropological Analysis of Blunt Force Trauma<\/em>. Springfield, IL: Charles C. Thomas Publisher, LTD.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Hefner, Joseph T., and Kandus C. Linde. 2018. <em>Atlas of Human Cranial <\/em><em>Macromorphoscopic<\/em><em> Traits<\/em>. San Diego: Academic Press.<\/p>\n<p class=\"import-Normal\">\u0130\u015fcan, M. Y., S. R. Loth, and R. K. Wright. 1984. \u201cAge Estimation from the Rib by Phase Analysis: White Males.\u201d <em>Journal of Forensic Sciences <\/em>29 (4): 1094\u20131104.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">\u0130\u015fcan, M. Y., S. R. Loth, and R. K. Wright. 1985. \u201cAge Estimation from the Rib by Phase Analysis: White Females.\u201d <em>Journal of Forensic Sciences <\/em>30 (3): 853\u2013863.Katz, Darryl, and Judy Myers Suchey. 1986. \u201cAge Determination of the Male Os Pubis.\u201d <em>American Journal of Physical Anthropology <\/em>69 (4): 427\u2013435.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Komar, Debra A., and Jane E. Buikstra. 2008. <em>Forensic Anthropology: Contemporary Theory and Practice<\/em>. New York: Oxford University Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Langley, Natalie R., Alice F. Gooding, and MariaTeresa Tersigni-Tarrant. 2017. \u201cAge Estimation Methods.\u201d In <em>Forensic Anthropology: A Comprehensive Introduction, Second Edition<\/em>, edited by Natalie R. Langley and MariaTeresa A. Tersigni-Tarrant, 175\u2013191. Boca Raton, FL: CRC Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Lovell, Nancy C. 1997. \u201cTrauma Analysis in Paleopathology.\u201d <em>Yearbook of Physical Anthropology<\/em> 104 (S25): 139\u2013170.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Native American Graves Protection and Repatriation Act (NAGPRA) 1990 (25 U.S. Code 3001 et seq.)<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">NIST (National Institute of Standards and Technology). N.d. \u201cThe Organization of Scientific Area Committees for Forensic Science.\u201d Accessed April 18, 2023. <a class=\"rId120\" href=\"https:\/\/www.nist.gov\/topics\/organization-scientific-area-committees-forensic-science\">https:\/\/www.nist.gov\/topics\/organization-scientific-area-committees-forensic-science<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ousley, Stephen. 1995. \u201cShould We Estimate Biological or Forensic Stature?\u201d <em>Journal of Forensic Sciences<\/em> 40(5): 768\u2013773.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Phenice, T. W. 1969. \u201cA Newly Developed Visual Method of Sexing the Os Pubis.\u201d <em>American Journal of Physical Anthropology<\/em> 30 (2): 297\u2013302.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Rose, Jerome C., Thomas J. Green, and Victoria D. Green. 1996. \u201cNAGPRA Is Forever: Osteology and the Repatriation of Skeletons.\u201d <em>Annual Review of Anthropology <\/em>25: 81\u2013103.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Schaefer, Maureen, Sue Black, and Louise Scheuer. <em>Juvenile Osteology: A Laboratory and Field Manua<\/em>l. 2009. San Diego: Elsevier Academic Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Schall, Jenna L., Tracy L. Rogers, and Jordan D. Deschamps-Braly. 2020. \u201cBreaking the Binary: The Identification of Trans-women in Forensic Anthropology.\u201d <em>Forensic Science International<\/em> 309: 110220. https:\/\/doi.org\/10.1016\/j.forsciint.2020.110220.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2010a. \u201cPersonal Identification.\u201d Last modified June 3, 2010. <a class=\"rId121\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_personal_identification.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_personal_identification.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2010b. \u201cSex Assessment.\u201d Last modified June 3, 2010. <a class=\"rId122\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_sex_assessment.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_sex_assessment.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2011. \u201cTrauma Analysis.\u201d Last modified May 27, 2011. <a class=\"rId123\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_trauma.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_trauma.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2012. \u201cStature Estimation.\u201d Last modified August 2, 2012. <a class=\"rId124\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_stature_estimation.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_stature_estimation.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Scientific Working Group for Forensic Anthropology (SWGANTH). 2013. \u201cAge Estimation.\u201d Last modified January 22, 2013. <a class=\"rId125\" href=\"https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_age_estimation.pdf\">https:\/\/www.nist.gov\/sites\/default\/files\/documents\/2018\/03\/13\/swganth_age_estimation.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Soler, Angela, and Jared S. Beatrice. 2018. \u201cExpanding the Role of Forensic Anthropology in Humanitarian Crisis: An Example from the USA-Mexico Border. In <em>Sociopolitics of Migrant Death and Repatriation: Perspectives from Forensic Science<\/em>, edited by Krista E. Latham and Alyson J. O\u2019Daniel, 115\u2013128. New York: Springer.<\/p>\n<p class=\"import-Normal\">Soler, Angela, Robin Reineke, Jared Beatrice, and Bruce E. Anderson. 2019. \u201cEtched in Bone: Embodied Suffering in the Remains of Undocumented Migrants.\u201d <em>In<\/em> <em>The Border and Its Bodies: The Embodiment of Risk along the U.S.-M\u00e9xico Line<\/em>, edited by Thomas E. Sheridan and Randall H. McGuire, 173\u2013207. Tucson: University of Arizona Press.<\/p>\n<p class=\"import-Normal\">Stull, Kyra E., Eric J. Bartelink, Alexandra R. Klales, Gregory E. Berg, Michael W. Kenyhercz, Erica N. L\u2019Abb\u00e9, Matthew C. Go, et al.. 2021. \u201cCommentary on: Bethard JD, DiGangi EA. Letter to the Editor\u2014Moving Beyond a Lost Cause: Forensic Anthropology and Ancestry Estimates in the United States. J Forensic Sci. 2020;65(5):1791\u20132. doi: 10.1111\/1556-4029.14513.\u201d <em>Journal of Forensic Sciences <\/em>66 (1): 417\u2013420.<\/p>\n<p class=\"import-Normal\">Tallman, Sean D., Caroline D. Kincer, and Eric D. Plemons. 2022. \u201cCentering Transgender Individuals in Forensic Anthropology and Expanding Binary Sex Estimation in Casework and Research.\u201d Special issue, \u201cDiversity and Inclusion,\u201d <em>Forensic Anthropology<\/em> 5 (2): 161\u2013180.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Tersigni-Tarrant, MariaTeresa A., and Natalie R. Langley. 2017. \u201cHuman Osteology.\u201d In <em>Forensic Anthropology: A Comprehensive Introduction, Second Edition<\/em>, edited by Natalie R. Langley and MariaTeresa A. Tersigni-Tarrant, 81\u2013109. Boca Raton, FL: CRC Press.<\/p>\n<p>Thurzo, A., Kosn\u00e1\u010dov\u00e1, H. S., Kurilov\u00e1, V., Kosme\u013e, S., Be\u0148u\u0161, R., Moravansk\u00fd, N., Kov\u00e1\u010d, P., Kuracinov\u00e1, K. M., Palkovi\u010d, M., &amp; Varga, I. (2021). Use of Advanced Artificial Intelligence in Forensic Medicine, Forensic Anthropology and Clinical Anatomy. Healthcare (Basel, Switzerland), 9(11), 1545. <a href=\"https:\/\/doi.org\/10.3390\/healthcare9111545\">https:\/\/doi.org\/10.3390\/healthcare9111545<\/a><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ubelaker, Douglas H. 2018. \u201cA History of Forensic Anthropology.\u201d Special issue, \u201cCentennial Anniversary Issue of AJPA,\u201d <em>American Journal of Physical Anthropology<\/em> 165 (4): 915\u2013923.<\/p>\n<p>Wankhade, T. D., Ingale, S. W., Mohite, P. M., &amp; Bankar, N. J. (2022). Artificial Intelligence in forensic medicine and toxicology: The future of forensic medicine. Cureus. <a href=\"https:\/\/doi.org\/10.7759\/cureus.28376\">https:\/\/doi.org\/10.7759\/cureus.28376<\/a><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">White, Tim D., and Pieter A. Folkens. 2005. <em>The Human Bone Manual<\/em>. Burlington, MA: Elsevier Academic Press.<\/p>\n<p class=\"import-Normal\">Winburn, Allysha P., and Bridget Algee-Hewitt. 2021. \u201cEvaluating Population Affinity Estimates in Forensic Anthropology: Insights from the Forensic Anthropology Database for Assessing Methods Accuracy (FADAMA).\u201d <em>Journal of Forensic Sciences<\/em> 66 (4): 1210\u20131219.<\/p>\n<p class=\"import-Normal\">Winburn, Allysha Powanda, Sarah Kiley Schoff, and Michael W. Warren. 2016. \u201cAssemblages of the Dead: Interpreting the Biocultural and Taphonomic Signature of Afro- Cuban Palo Practice in Florida.\u201d <em>Journal of African Diaspora Archaeology and Heritage <\/em>5 (1): 1\u201337.<\/p>\n<\/div>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1114\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1114\"><div tabindex=\"-1\"><p>The act of searching for food.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_382_1784\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_382_1784\"><div tabindex=\"-1\"><p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.791999816894531pt;margin-right: 0pt;text-indent: 0pt\">Joylin Namie, Ph.D., Truckee Meadows Community College<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\"><span style=\"color: #ffffff\">Learning Objectives<\/span><\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li>Identify and describe the major developments in scientific thought that led to the discovery of evolutionary processes.<\/li>\n<li>Explain how natural selection works and results in evolutionary change over time.<\/li>\n<li>Explain what is meant by the \u201cModern Synthesis\u201d and its impacts on evolutionary thought.<\/li>\n<li>Discuss the teaching of human evolution in the U.S. and abroad.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<h2 class=\"__UNKNOWN__\">The Beginnings of Evolutionary Thinking<\/h2>\n<div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.627006530761719pt;margin-right: 3.917236328125pt;text-indent: 0pt\">Throughout our evolutionary history, humans have developed an understanding of the natural world as they interacted with and extracted resources from it. To survive, our earliest ancestors possessed an understanding of the physical environment, including weather patterns, animal behavior, edible and medicinal plants, locations of water, and seasonal cycles. Many ancient cultures, including those of the Americas (Dunbar-Ortiz 2014), Mesopotamia, and Egypt, left writings, hieroglyphics, and stories passed down through oral tradition detailing their understanding of the natural environment, human and zoological anatomy, botany, and medical practices (Moore 1993).<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 1.209999084472656pt;margin-right: 11.6515502929688pt;text-indent: 0pt\">There are also over 2,000 years of organized thought and writing regarding <strong>evolution<\/strong>, including contributions from Greek, Roman, and Islamic scholars. Three examples of note are included here. The Greek philosopher Aristotle (384\u2013322 BCE) studied the natural world, publishing several volumes on animals based on systematic observations, rather than attributing what he observed to divine intervention, as his contemporaries were doing (Figure 2.1). Aristotle\u2019s system for the biological classification of nearly 500 species of animals was based on his own observations and dissections, interviews with specialists such as beekeepers and fishermen, and accounts of travelers. His nine book <em>History of Animals<\/em>, published in the 4th century BC (n.d.), was one of the first zoological taxonomies ever created. Aristotle\u2019s primary contribution to the classification of biological species was to recognize that natural groups are based on structure, physiology, mode of reproduction, and behavior (Moore 1993, 39).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 1.243003845214844pt;margin-right: 31.8323364257812pt;text-indent: 0pt\"><img class=\"aligncenter\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/03\/image3-5.jpg\" alt=\"Large orange octopus on ocean floor.\" width=\"240\" height=\"321\" \/><\/p>\n<figure style=\"width: 338px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image12-4.jpg\" alt=\"Elephant half-submerged in a body of water with a ferry of human watchers behind.\" width=\"338\" height=\"231\" \/><figcaption class=\"wp-caption-text\">Figure 2.1a-b: Aristotle was the first to publish that a. octopuses can change their colors when disturbed and b. elephants use their trunks as a snorkel when crossing deep water. Credit: a. <a href=\"https:\/\/en.wikipedia.org\/wiki\/File:Octopus_macropus.jpg\">Octopus macropus<\/a> by <a href=\"https:\/\/subnormali-team.blogspot.com\/2006_12_01_archive.html\">SUBnormali Team<\/a> (originally from <a href=\"https:\/\/it.wikipedia.org\/wiki\/Utente:Yoruno\">Yoruno<\/a>) is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/#:~:text=You%20are%20free%20to%3A,for%20any%20purpose%2C%20even%20commercially.\">CC-BY-SA 3.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Elephant_swimming,_Botswana_(cropped).jpg\">Elephant swimming, Botswana (cropped)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/8721758@N06\">Jorge L\u00e1scar<\/a> from Australia (uploaded by <a href=\"https:\/\/www.flickr.com\/photos\/29050464@N06\/\">Peter D. Tillman<\/a>) is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/deed.en\">CC BY 2.0 License<\/a>.<\/figcaption><\/figure>\n<figure style=\"width: 263px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image20-3.png\" alt=\"Rows of organisms, with plants and animals at the bottom and humans, angels, and God at top.\" width=\"263\" height=\"379\" \/><figcaption class=\"wp-caption-text\">Figure 2.2: The Great Chain of Being by Didacus Valades. Credit: <a href=\"https:\/\/commons.wikimedia.org\/w\/index.php?curid=1688250\">Great Chain of Being<\/a> by Didacus Valades (Diego Valades 1579) and photographed by Rhetorica Christiana (via <a href=\"https:\/\/archive.org\/details\/rhetoricachristi00vala\/page\/n259\/mode\/2up\">Getty Research<\/a>) is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 11.6515502929688pt;text-indent: 0pt\">Aristotle\u2019s <em>History of Animals<\/em> also placed animals in a hierarchy, ranking animals above plants due to what he claimed were their abilities to sense the world around them and to move. He also graded animals according to their modes of reproduction. Those giving birth to live young were placed above those who laid eggs. Warm-blooded animals ranked above invertebrates. This concept of \u201chigher\u201d and \u201clower\u201d organisms was expanded upon by scholars in the Medieval period to form the <em>Scala Naturae<\/em> (Latin for \u201cladder of being\u201d). This \u201cGreat Chain of Being,\u201d depicting a hierarchy of beings with God at the top and minerals at the bottom (Figure 2.2), was thought by medieval Christians to have been decreed by God; in this Great Chain, humans were placed closer to God than other species. Aristotle\u2019s works were rediscovered by Islamic scholars in the ninth century and translated into Arabic, Syriac, Persian, and later into Latin, becoming part of university curriculum in 13th-century Europe (Lindberg 1992), allowing Aristotle\u2019s works and ideas to influence other thinkers for 2,000 years.<\/p>\n<figure style=\"width: 251px\" class=\"wp-caption alignright\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image8-5.jpg\" alt=\"A person leading a giraffe on a leash, with text written in Arabic below.\" width=\"251\" height=\"375\" \/><figcaption class=\"wp-caption-text\">Figure 2.3: An image from Kit\u0101b al-\u1e25ayaw\u0101n (Book of the Animals) by Al-Jahiz. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Al-Jahiz.jpg\">Al-Jahiz<\/a> by Al-Jahiz [in <a href=\"https:\/\/themuslimtimes.info\/2017\/02\/25\/al-jahizs-book-of-animals-the-transcendent-value-of-disgust\/\">Kit\u0101b al-\u1e25ayaw\u0101n<\/a> (Book of the Animals), 15th century] is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 12.08837890625pt;text-indent: 0pt\">Science also owes a debt to many Arabic scholars. One such Islamic scholar and writer, who built upon the ideas of Aristotle, was Ab\u016b \u02bfUthman \u02bfAmr ibn Ba\u1e25r al-Kin\u0101n\u012b al-Ba\u1e63r\u012b \/ al-J\u0101\u1e25i\u1e93, known as Al-Jahiz (776\u2013868 CE), who authored over 200 books (El-Zaher 2018; Figure 2.3). His most well-known work was the seven-volume <em>Kitab al-Hayawan<\/em> or <em>Book of Animals<\/em>, in which he described over 350 species in zoological detail. Importantly, Al-Jahiz introduced the idea and mechanisms of biological evolution 1,000 years before Darwin proposed the concept of <strong>natural selection<\/strong> in 1859 (Love 2020). For instance, Al-Jahiz wrote about the struggle for existence, the transformation of species over time, and environmental factors that influence the process, all ideas that were later espoused by western European scientists in the 19th century. Al-Jahiz wrote:<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.891006pt;margin-right: 12.0884pt;text-indent: 0.648994pt;text-align: left;padding-left: 40px\">Animals engage in a struggle for existing, and for resources, to avoid being eaten, and to breed. Environmental factors influence organisms to develop new characteristics to ensure survival, thus transforming them into new species. Animals that survive to breed can pass on their successful characteristics to their offspring. [Masood 2009]<\/p>\n<figure style=\"width: 335px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image9-7.png\" alt=\"A person with a full beard and turban looks into the distance.\" width=\"335\" height=\"389\" \/><figcaption class=\"wp-caption-text\">Figure 2.4: Drawing of Ibn al-Haytham. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Ibn_al-Haytham.png\">Ibn al-Haytham<\/a> by Sopianwar is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.891006469726562pt;margin-right: 12.08837890625pt;text-indent: 0.648994445800781pt\">Another important early Islamic scientist is Ibn al-Haytham (965\u20131040), a 10th-century Islamic scholar who contributed a great deal to our understanding of optics and how human vision works (Figure 2.4; Lindberg 1992, 177\u2013180). Born in what is now Iraq, al-Haytham was a scholar of many disciplines, including mathematics, physics, mechanics, astronomy, philosophy, and medicine. He authored some 200 books in his lifetime and was an expert on Aristotle\u2019s natural philosophy, logic, and metaphysics. In relation to evolution, al-Haytham\u2019s methodology of investigation\u2014specifically, using experiments to verify theory\u2014is similar to what later became known as the modern scientific method. He is most famous for discovering the laws of reflection and refraction over 1,000 years ago and inventing the camera obscura, which was incredibly important for the eventual development of photography. His work is credited for its influence on astronomy, mathematics, and optics, inspiring Galileo, Johannes Kepler, and Sir Isaac Newton (Tasci 2020). As an inspirational scientific figure, al-Haytham was celebrated in 2016 by UNESCO as a trailblazer and the founder of modern optics (Figure 2.5). An International Year of Light was named in his honor and a scholarly conference on his contributions was held to coincide with the 1,000th anniversary of the publication of his <em>Kit\u0101b al-Man\u0101\u1e93ir<\/em> (Book of Optics; UNESCO.org 2015).<\/p>\n<figure style=\"width: 239px\" class=\"wp-caption alignright\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image16-2.jpg\" alt=\"Labeled diagram of the eye and optic nerves.\" width=\"239\" height=\"399\" \/><figcaption class=\"wp-caption-text\">Figure 2.5: Diagram of the Human Eye by Ibn al-Haytham. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:I.A._Haitham,_Diagram_of_the_eye,_16th_century_Wellcome_L0011969.jpg\">Diagram of the eye<\/a> by Ibn Al [Alhazen] Haitham (16th Century) has been modified (cropped) and is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/legalcode\">CC BY 4.0 License<\/a>. This image is available from <a href=\"https:\/\/wellcomeimages.org\/\">Wellcome Images<\/a> 3044 (under the photo number L0011969).<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0.891006469726562pt;margin-right: 12.08837890625pt;text-indent: 0.648994445800781pt\">The writings of these Islamic scholars as well as similar scientific texts from other cultures were unknown to or unacknowledged by Western scientists until recently. Fortunately, many science teachers are now incorporating this content into their classes (Love 2020).<\/p>\n<h2 class=\"import-Normal\" style=\"background-color: transparent;margin-left: 1.364006042480469pt;margin-right: 0pt;text-indent: 0pt\">Western European Evolutionary Thought<\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.836006164550781pt;margin-right: 0.0257568359375pt;text-indent: 0pt\">Although there have been many different scientific traditions throughout world history, a new global discourse around science emerged in Western Europe in the 19th century. Europeans pointed to the continuing expansion of their colonial power, as well as their military and technological success, as evidence of the efficacy of Western science, which came to dominate on a global scale (Elshakry 2010). The movement toward a global science centered in Western Europe began with formulation of the <strong>Scientific Method<\/strong>.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0.0257568359375pt;text-indent: 0pt\">The Scientific Method was first codified by Francis Bacon (1561\u20131626), an English politician who was likely influenced by the methods of inquiry established by Ibn al-Haytham centuries prior (Tbakhi and Amr 2007). Bacon has been called the founder of <strong>empiricism<\/strong> for proposing a system for weighing the truthfulness of knowledge based solely on inductive reasoning and careful observations of natural phenomena. Ironically, he died as a result of trying to scientifically observe the effects of cold on the putrefaction of meat. On a journey out of London, he purchased a chicken and stuffed it with snow for observation, catching a chill in the process. One week later, he died of bronchitis (Urbach, Quinton, and Lea 2023).<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0.0257568359375pt;text-indent: 0pt\">The second important development with regard to evolution was the concept of a <strong>species<\/strong>. John Ray (1627\u20131705), an English parson and naturalist, was the first person to publish a biological definition of species in his <em>Historia Plantarum<\/em> (<em>History of Plants),<\/em> a three volume work published in 1686, 1688, and 1704<em>. <\/em>Ray defined a <em>species<\/em> as a group of morphologically similar organisms arising from a common ancestor. However, we now define a species as a group of similar organisms capable of producing fertile offspring. In keeping with the scientific method, Ray classified plants according to similarities and differences that emerged from observation. He claimed that any seed from the same plant was the same species, even if it had slightly different traits.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.836006164550781pt;margin-right: 4.403076171875pt;text-indent: 0pt\">The modern period of biological classification began with the work of Carl von Linne (\u201cCarolus Linnaeus\u201d) (1707\u20131778), a Swedish scientist who laid the foundations for the modern scheme of taxonomy used today. He established the system of <strong>binomial nomenclature<\/strong>, in which a species of animal or plant receives a name consisting of two terms: the first term identifies the genus to which it belongs and the second term identifies the species. His original <em>Systema<\/em> <em>Naturae<\/em>, published in 1736, went through several editions. By the tenth edition in 1758, mammals incorporated primates, including apes and humans, and the term <em>Homo sapiens <\/em>was introduced to signify the latter (Paterlini 2007).<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.495002746582031pt;margin-right: 4.97509765625pt;text-indent: 0pt\">Georges-Louis Leclerc, Comte de Buffon (1707\u20131788), was a prominent French naturalist whose work influenced prominent scientists in the second half of the 18th century. Buffon's idea that species change over time became a cornerstone of modern evolutionary theory. His technique of comparing similar structures across different species, called <strong>comparative anatomy<\/strong>, is still in use today in the study of evolution. He published 36 volumes of <em>Histoire<\/em> <em>Naturelle<\/em> during his lifetime and heavily influenced two prominent French thinkers who were to have significant impacts on our understanding of evolution, Georges Cuvier and Jean-Baptiste Lamarck.<\/p>\n<figure style=\"width: 455px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image23-2.jpg\" alt=\"Historic painting of person with short wavy hair next to drawing of a mastodon skeleton.\" width=\"455\" height=\"302\" \/><figcaption class=\"wp-caption-text\">Figure 2.6: Cuvier with one of his drawings of a fossil quadruped. Credit: Cuvier and a fossil quadruped original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) is a collective work under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Includes <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Georges_Cuvier_3.jpg\">Georges Cuvier 3<\/a> by <a href=\"https:\/\/en.wikipedia.org\/wiki\/Fran%C3%A7ois-Andr%C3%A9_Vincent\">Fran\u00e7ois-Andr\u00e9 Vincent<\/a> (artist), <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>; <a href=\"https:\/\/freesvg.org\/mammoth-skeleton\">Mammoth skeleton<\/a> in <a href=\"https:\/\/freesvg.org\/by\/OpenClipart\">OpenClipart<\/a>, <a href=\"https:\/\/creativecommons.org\/share-your-work\/public-domain\/cc0\/\">public domain (CC0)<\/a>.]<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.594001770019531pt;margin-right: 1.4913330078125pt;text-indent: 0pt\">Georges Cuvier (1769\u20131832) was a paleontologist and comparative anatomist (Figure 2.6). One of his first major contributions to the field of evolution was proof that some species had become <strong>extinct <\/strong>through detailed and comprehensive analyses of large fossil quadrupeds (Moore 1993, 111). The idea of extinction was not new, but it was challenging to demonstrate if a fossil species was truly extinct or still had living relatives elsewhere. It was also challenging in that it ran counter to religious beliefs of the time. The Bible\u2019s Book of Genesis was interpreted as saying that all species had been created by God in the seven days it took to create the world and that all created species have survived to this day. Extinction was interpreted as implying imperfection, suggesting God\u2019s work was flawed. Also, given that the Earth was calculated to have been created in 4004 B.C.E., based on biblical genealogies, there would not have been enough time for species to disappear (Moore 1993, 112).<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 1.4913330078125pt;text-indent: 0pt\">Cuvier was so knowledgeable in this field that he became famous for his ability to reconstruct what an extinct animal looked like from fragmentary remains. He demonstrated that fossil mammoths differed from similar living creatures, such as elephants. His many examples of fossils telling the stories of animals that lived and then disappeared were taken as incontrovertible proof of extinctions (PBS 2001). Where Cuvier went awry was his hypothesis of how extinction worked and its causes. As part of his study of comparative anatomy, Cuvier made observations of stratified layers of rock, or sediment, each containing different species. From this, he drew conclusions that species were \u201cfixed\u201d and did not evolve, but then went extinct, and that different assemblages of fossils occurred at different times in the past, as evidenced by the sedimentary layers (Moore 1993, 118). Cuvier explained this through a theory of <strong>catastrophism<\/strong>, which stated that successive catastrophic deluges (akin to Biblical floods) swept over parts of the Earth periodically, exterminating all life. When the waters receded from a particular region, lifeforms from unaffected regions would repopulate the areas that were destroyed, giving rise to a new layer of species that looked different from the layer below it. This theory implied that species were fixed in place and did not evolve and that the Earth was young. In fact, Cuvier postulated that the last catastrophe was a deluge he believed occurred five to six thousand years ago, paving the way for the advent of humans (Moore 1993, 118). Cuvier\u2019s catastrophism became part of an ongoing and vociferous debate between two schools of geology. The catastrophists believed the present state of the earth was the consequence of a series of violent catastrophes of short duration, while the uniformitarians thought it was the result of slow acting geological forces that continue to shape the earth.<\/p>\n<figure style=\"width: 379px\" class=\"wp-caption alignright\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image13-2.jpg\" alt=\"Horizontal layers of rock rest on vertical layers of rock.\" width=\"379\" height=\"303\" \/><figcaption class=\"wp-caption-text\">Figure 2.7: Siccar Point, Aberdeen, UK. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Siccar_Point.jpg\">Siccar Point<\/a> by <a href=\"https:\/\/www.geograph.org.uk\/profile\/139\">Anne Burgess<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/legalcode\">CC BY-SA 2.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.495002746582031pt;margin-right: 1.47552490234375pt;text-indent: 0pt\">James Hutton (1726\u20131797) was one prominent proponent of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_830\">uniformitarianism<\/a><\/strong>. Based on evidence he found at sites in his native Scotland, Hutton argued that the Earth was much older than previously thought. Examining the geology of Siccar Point, a cliff site on the eastern coast of Scotland (Figure 2.7), Hutton concluded that the intersection of the vertical and horizontal rocks represented a gap in time of many millions of years, during which the lower rocks had been deformed and eroded before the upper layers were deposited on top. From this, Hutton argued sediments are deposited primarily in the oceans, where they become strata, or layers of sedimentary rock. Volcanic action uplifts these strata to form mountains, which are then subject to erosion from rain, rivers, and wind, returning sediment to the oceans (Moore 1993, 121). Hutton\u2019s <em>Theory of the Earth <\/em>(1788) demanded vast periods of time (known as \u201cdeep time\u201d) for such slow-working forces to shape the earth. At the time, he was heavily criticized for this view, as it contradicted the biblical version of the history of creation.<\/p>\n<figure style=\"width: 391px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image18-3.jpg\" alt=\"A cross-section of a volcanic eruption showing different types of rock that make up the volcano.\" width=\"391\" height=\"249\" \/><figcaption class=\"wp-caption-text\">Figure 2.8: The frontispiece from Charles Lyell's Principles of Geology (2nd American edition, 1857), showing the origins of different rock types. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Lyell_Principles_frontispiece.jpg\">Lyell Principles frontispiece<\/a> by Charles Lyell (Principles of Geology, 2nd American edition, 1857) is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.495002746582031pt;margin-right: 1.47552490234375pt;text-indent: 0pt\">Another Scotsman, who was to become a highly influential geologist and a close friend of Darwin, was Charles Lyell (1797\u20131875). Lyell was originally a lawyer who began his studies of Geology at Oxford under the tutelage of catastrophist William Buckland, from whom he diverged when Buckland tried to find physical evidence of Noah\u2019s flood from the Christian Bible. Lyell was instead intent on establishing geology as a science based on observation. Building upon Hutton\u2019s ideas (published 50 years earlier), Lyell traveled throughout Europe, documenting evidence of uniformitarianism. During his travels, he cataloged evidence of sea level rise and fall and of volcanoes positioned atop much older rocks. He also found evidence of valleys formed through erosion, mountains resulting from earthquakes, and volcanic eruptions that had been witnessed or documented in the past (University of California Berkeley Museum of Paleontology n.d.). Lyell also espoused the principle that \u201crocks and strata (layers of rock) increase in age the further down they are in a geological sequence. Barring obvious upheavals or other evidence of disturbance, the same principle must apply to any fossils contained within the rock. The lower down in a sequence of rocks a fossil is, the older it is likely to be (Wood 2005, 12).\u201d<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0.495002746582031pt;margin-right: 1.47552490234375pt;text-indent: 0pt\">Lyell published the first edition of his three-volume <em>Principles of Geology <\/em>in 1830\u20131833 (Figure 2.8). It established geology as a science, underwent constant revisions as new scientific evidence was discovered, and was published in 12 editions during Lyell\u2019s lifetime. In it, he espoused the key concept of uniformitarianism\u2014that \"the present is the key to the past.\u201d What this meant was that geological remains from the distant past can be explained by reference to geological processes now in operation and directly observable.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 1.83465576171875pt;text-indent: 0pt\">Jean-Baptiste Lamarck (1744\u20131829) was the first Western scientist to propose a mechanism explaining why and how traits changed in species over time, as well as to recognize the importance of the physical environment in acting on and shaping physical characteristics. Lamarck\u2019s view of how and why species changed through time, known as the \u201cTheory of Inheritance of Acquired Characteristics,\u201d was first presented in the introductory lecture to students in his invertebrate zoology class at the Museum of Natural History in Paris in 1802 (Burkhardt 2013). It was based on the idea that as animals adapted to their environments through the use and disuse of characteristics, their adaptations were passed on to their offspring through reproduction (Figure 2.9). Lamarck was right about the environment having an influence on characteristics of species, as well as about variations being passed on through reproduction. He simply had the mechanism wrong.<\/p>\n<p><img class=\"aligncenter\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image14-5.png\" alt=\"Giraffes with necks of different heights reach to eat leaves.\" width=\"419\" height=\"244\" \/><\/p>\n<figure style=\"width: 404px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image11-7.png\" alt=\"Illustration of three giraffes with necks of different heights.\" width=\"404\" height=\"391\" \/><figcaption class=\"wp-caption-text\">Figure 2.9a-b: Inheritance of Acquired Characteristics and Natural Selection. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Lamarckian Evolution (Figure 4.2A and 4.2B)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 1.86676025390625pt;text-indent: 0pt\">Lamarck\u2019s theory involved a three-step process. Step one involves an animal experiencing a radical change in its environment. Step two is the animal (either individual or species) responding with a new kind of behavior. Step three is how the behavioral change results in morphological (meaning physical) changes to the animal that are successfully passed on to subsequent generations (Ward 2018, 8). Lamarck\u2019s most famous example was the proposition that giraffes actively stretched their necks to reach leaves on tall trees to eat. Over their lifetimes, the continuation of this habit resulted in gradual lengthening of the neck. These longer necks were then passed on to their offspring. Lamarck's theory was disproved when evolutionary biologist August Weismann published the results of an experiment involving mice (Figure 2.10). Weismann amputated the tails of 68 mice and then successively bred five generations of them, removing the tails of all offspring in each generation, eventually producing 901 mice, all of whom had perfectly healthy long tails in spite of having parents whose tails were missing (Weismann 1889).<\/p>\n<figure style=\"width: 551px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image6-5.png\" alt=\"Mice with cut-off ails breed healthy offspring with full length tails.\" width=\"551\" height=\"385\" \/><figcaption class=\"wp-caption-text\">Figure 2.10: Weismann\u2019s mouse-tail experiment showing that offspring do not inherit traits that the parents acquired during their lifetimes. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Weismann\u2019s mouse-tail experiment (Figure 4.3)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.527999877929688pt;margin-right: 0.20892333984375pt;text-indent: 0pt\">How giraffes actually ended up with long necks is a different story. In an environment where the food supply is higher off the ground, and perhaps less available to competing species, giraffes who happened to have slightly longer necks (due to random individual variation and genetic mutation) would be more likely to survive. These giraffes would then be able to reproduce, passing along the slight variation in neck length that would allow their offspring to do the same. Over time, individuals with longer necks would be overrepresented in the population, and neck lengths overall would increase among giraffes. Unfortunately, Lamarck\u2019s ideas challenged the scientific establishment of the time and were rejected. He was discredited and harassed \u201cto the point of loss of money, reputation, and then health\u201d (Ward 2018, 9).<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.527999877929688pt;margin-right: 0.4715576171875pt;text-indent: 0pt\">The final piece in the evolutionary puzzle leading up to the theory of natural selection was put forth by Thomas Malthus (1766\u20131834), who published <em>A<\/em><em>n Essay on Population <\/em>in 1798. Malthus lived in England during the time of the Industrial Revolution. It was a time of great poverty and misery when many people migrated from the countryside to squalid, disease-ridden cities to work extremely long hours in dangerous conditions in factories, coal mines, and other industrial workplaces. Birth rates were high and starvation and disease were rampant. Malthus struggled to explain why. His answer was basically the idea of <strong>carrying capacity<\/strong>, an ecological concept still in use today. Malthus suggested the rate of population growth exceeded the rate of increase of the human food supply. In other words, people were outgrowing the available food crops. He also suggested that populations of animals and plants were naturally constrained by the food supply, resulting in reductions in population in times of scarcity, \u201crestraining them within the prescribed bounds\u201d (Moore 1993, 147). But, despite significant challenges, some individuals always survived. This was the key to later understandings of evolutionary change in species over time.<\/p>\n<h2 class=\"import-Normal\" style=\"background-color: transparent;margin-left: 1.331001281738281pt;margin-right: 0pt;text-indent: 0pt\">The Journey to Natural Selection<\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.527999877929688pt;margin-right: 7.171630859375pt;text-indent: 0pt\">In Western European thought, the individual most closely associated with evolution is Charles Darwin (1809\u20131882). However, as one can see from the individuals and ideas presented in the prior section, he was not the first person to explore evolution and how it might work. In fact, Darwin built upon and synthesized many of the ideas\u2014from geology to biology, ecology, and economy\u2014discussed above. He was simply in the right place at the right time. If he had not worked out his ideas when he did, someone else would have. As a matter of fact, as noted below, someone else did, forcing Darwin to publicly reveal his theory.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 7.171630859375pt;text-indent: 0pt\">Darwin continued his observations and experiments during his formal education, culminating in his graduation from Cambridge in 1831, at which point he was invited to become a gentleman naturalist for a British Royal Navy surveying mission of the globe aboard the H.M.S. <em>Beagle<\/em>. It is worth noting that Darwin was only 22 years old and the captain\u2019s third choice for the position (Costa 2017), but he proved extremely curious and methodical. The mission departed in December of 1831 and returned five years later (Figure 2.11). During this time, Darwin produced copious notebooks, observations, drawings, and reflections on the natural phenomena he encountered and the experiments he performed.<\/p>\n<figure style=\"width: 780px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image22-5.png\" alt=\"The voyage of the Beagle throughout the world.\" width=\"780\" height=\"329\" \/><figcaption class=\"wp-caption-text\">Figure 2.11: Map of the voyage of the H.M.S. Beagle. <a href=\"https:\/\/docs.google.com\/document\/d\/1VUDKMBJYS_jNONjLxT04jQN0_z9Ua50BRN6auGSHUuU\/edit\" target=\"_blank\" rel=\"noopener\">A full text description of this image is available.<\/a> Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Voyage_of_the_Beagle-de.svg\">Voyage of the Beagle-de<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Succu\">Succu<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 3.4952392578125pt;text-indent: 0pt\">Discussing all of Darwin\u2019s work aboard the <em>Beagle <\/em>is beyond the scope of this chapter, but his primary interests were in cataloging new varieties of plant and animal life and examining the geology of the places the ship made landfall. Part of Darwin\u2019s success with regard to both ventures was due to his extreme seasickness, which began before the ship even left Plymouth Harbor. It never let up, encouraging Darwin to go ashore at every available opportunity. \u201cIn fact, of the nearly five years of the voyage, Darwin was actually on board the ship for just a year and a half altogether\u201d (Costa 2017, 18).<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.527999877929688pt;margin-right: 4.93707275390625pt;text-indent: 0pt\">During the voyage, the young Darwin tried to make sense of what he saw through the lens of the scientific paradigms he held when he left England, but he continually made observations that challenged these paradigms. For example, while the <em>Beagle<\/em> crewmen were charting the coast of Argentina, Darwin conducted fieldwork on land. There he observed species that were new to him, like armadillos. He also collected fossils, including those of extinct armadillos. Meaning, he had found both <strong>extant <\/strong>and extinct members of the same species in the same geographic location, which challenged the theory of catastrophism put forth by Cuvier, who argued that each variant of an animal, living or extinct, was its own distinct species (Moore 1993, 144). Darwin also observed geographic variation in the same species all along the east coast of South America, from Brazil to the southern tip of Argentina. He noted that some species were found in multiple localities and differed from place to place. Those living closer to each other exhibited only slight variations, while those living further apart might be cataloged as entirely different species if one did not know better.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.693000793457031pt;margin-right: 14.136962890625pt;text-indent: 0pt\">He made similar observations in the Galapagos Islands located off the northwest coast of Ecuador, with regard to giant tortoises and finches (Figure 2.12). A local resident of the islands explained to Darwin that each island had its own variety of tortoise and that locals could discern which island a tortoise came from simply by looking at it. Darwin noted other such examples in both plants and animals, meaning geographic variation was occurring on separate, neighboring islands.<\/p>\n<figure style=\"width: 684px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image15-4.jpg\" alt=\"Hood Island tortoises have saddle-backed shells; Isabella Island, dome-shaped; Pinta Island, intermediate.\" width=\"684\" height=\"528\" \/><figcaption class=\"wp-caption-text\">Figure 2.12: Variation in giant tortoises in the Galapagos Islands. Credit: Giant Tortoises of the Galapagos Islands original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Mary Nelson and Katie Nelson is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0.67529296875pt;text-indent: 0pt\">Prevailing views of time argued that variations in living species, and even the fossil armadillos and the living armadillos, were the result of separate creation events. According to this view, each variation, no matter how slight, was a different species. Challenging these ideas would mean challenging not only catastrophism, but the <strong>Fixity of Species<\/strong> and other well-accepted ideas of the time. Darwin was aware that he was a young, unestablished naturalist. He was also aware of the ruin that befell Lamarck when his theories were rejected. Lastly, Lyell, who was a good friend of Darwin\u2019s, rejected evolution altogether. It is no wonder that Darwin published a great deal about the geological and fossil data he collected when he returned from the voyage, but not his early hypotheses about evolution.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.627006530761719pt;margin-right: 0pt;text-indent: 0pt\">Upon Darwin\u2019s return to England, it took another twenty years of data collection and experimentation before he was ready to share his conclusions about evolution. Much of this work was conducted at Down House, his home of forty years, where he performed all sorts of experiments that laid the groundwork for his ideas about evolution. Darwin\u2019s home was his laboratory, and he engaged the help of his children, neighbors, friends, and servants in collecting, dissecting, and experimenting. At one point in the 1850s, sheets of moistened paper covered with frogs eggs lined the hallways of the house, while flocks of sixteen different pigeon breeds cooed outside, glass jars filled with salt water and floating seeds filled the cellar, and the smell of dissected pigeon skeletons pervaded the air inside the house. There were also ongoing experiments in the yard, including piles of dissected flowers, beekeeping, and fenced-off plots of land where seedlings were under study. Darwin was a keen experimental scientist, observer, and a prolific writer and presenter of scientific papers. He regarded his work as \u201cone long argument\u201d that never really ended. In fact, Darwin published ten books after <em>On the Origin of Species<\/em>, addressing such far-ranging topics as animal behavior, orchids, and domestication, among others (Costa 2017).<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.627006530761719pt;margin-right: 0pt;text-indent: 0pt\">Darwin may not have published <em>Origins<\/em> in 1859 had it not been for receiving a paper in June of 1858 from Alfred Russel Wallace, an English naturalist working in Malaysia, espousing the same ideas. Wallace had sent the paper to Darwin asking if it was worthy of publication and requesting he forward it to Lyell and the English botanist, Joseph Hooker. Darwin wrote to Lyell and Hooker about Wallace\u2019s paper, entitled <em>On the Tendency of Varieties to Depart Indefinitely from the Original Type<\/em>. In recognition that both Wallace and Darwin had arrived at the same discovery, a \u201cjoint\u201d paper composed of four parts (none of them actually coauthored) was read to the Linnaean Society by Lyell, then secretary of the Society, on July 1, 1858, and published on August 20. Darwin published <em>On the Origin of Species <\/em>15 months later. (The original composite paper read before the Linnaean society is available to read for free from the Alfred Russell Wallace Website, on the <a href=\"https:\/\/wallacefund.myspecies.info\/content\/1858-darwin-wallace-paper\">1858 Darwin-Wallace paper<\/a> page.)<\/p>\n<h3 class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.726005554199219pt;margin-right: 0pt;text-indent: 0pt\"><strong>The Mechanism of Natural Selection <\/strong><\/h3>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.693000793457031pt;margin-right: 4.15203857421875pt;text-indent: 0pt\">Let us take a moment here to explore the mechanism of natural selection in more detail. Before we begin, it is important to recognize that Darwin defined evolution as descent with modification, by which he meant that species share a common ancestor yet change over time, giving rise to new species. Descent with modification refers to the fact that offspring from two parents look different from each of their parents, and from each other, meaning they descend with slight differences (\u201cmodifications\u201d). If you have ever observed a litter of puppies or a field of flowers and stopped to examine each individual closely, you have seen that each differs from the next, and none look exactly like their parents. These variations are random, not specific, and may or may not be present in the following generations.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.693000793457031pt;margin-right: 4.15203857421875pt;text-indent: 0pt\">Darwin struggled to explain why some slight differences were preserved over time, while others were not. He turned to what he knew of animal breeding (<strong>artificial selection<\/strong>) for an explanation (Richards 1998). Darwin bred different breeds of pigeons at Down House, carefully documenting phenotypic differences across generations, including slight anatomical variations he observed through dissection. He also grew and crossbred species of flowers and dissected those too. Darwin was also very fond of hunting and of hunting dogs. In an early draft of his theory on speciation, he used greyhounds as an example of adaptation and selection, \u201cnoting how its every bone and muscle, instinct and habit, were fitted to run down hare (rabbits) (University of Cambridge n.d.).\u201d In each case of plant and animal breeding Darwin observed, he noted that humans were selecting variants in each generation that had characteristics humans desired (i.e., sweetness of fruits, colors of flowers, fur type and color of animals). Breeders then continually bred plants and animals with the desired variants, over and over again. These small changes added up over time to create new species of plants and breeds of animals. Darwin also noted that artificial selection does not necessarily render plants or animals better adapted to their original environments. The characteristics humans desire often result in plants less likely to survive in the wild and animals more likely to suffer from certain behavioral or health problems. One has only to examine high rates of hip dysplasia in several modern breeds of dogs to observe what Darwin was referring to.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.693000793457031pt;margin-right: 4.15203857421875pt;text-indent: 0pt\">From his studies of artificial selection, Darwin drew the conclusion that nature also acts upon variations among members of the same species. Instead of human intervention, the forces of nature, such as heat, cold, predation, disease, and now climate change, determine which offspring, with which variants, survive and reproduce. These individuals then pass down these favorable variants to their own offspring. In this way, nature selects for traits that are beneficial within a particular environment and selects against traits that are disadvantageous within a particular environment. Over many generations, populations of a species become more and more adapted (or, in evolutionary terms, \u201cfit\u201d) for their specific environments. Darwin named this process natural selection.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.693000793457031pt;margin-right: 4.15203857421875pt;text-indent: 0pt\">This theory explained the variations in tortoises Darwin had observed years earlier in the Galapagos Islands (see Figure 2.12). Tortoises who lived on larger islands with lush vegetation to feed on were larger than those on smaller islands. They also had shorter necks and dome-shaped shells as their food was close to the ground. Tortoises on smaller, drier islands fed on cacti, which grew much taller. These tortoises had longer necks, longer front legs, and saddle-shaped shells, which allowed them to successfully stretch to reach the edible cactus pads that grew on the tops of the plants. How did these observable differences in the two tortoise populations emerge? Darwin would argue that, over time, small, random variations in the tortoises were differentially selected for by the distinct natural environments on different islands.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.693000793457031pt;margin-right: 4.15203857421875pt;text-indent: 0pt\">In addition to the biogeographical evidence Darwin offered from his research aboard the <em>Beagle<\/em>, as well as the evidence he documented from the artificial selection of plants and animals, he also relied, where possible, on fossil evidence. One example, mentioned above, were the fossil findings of extinct armadillos in Argentina in the same locations as living armadillos. Unfortunately, as Darwin himself noted, the geological record was incomplete, most often missing the transitional fossil forms that bridge extinct and living species. That issue has since been resolved with scientific research in geochronology and paleontology, among other fields. It is now well-established that life is far more ancient than was believed in Darwin\u2019s time and that these ancient forms of life were the ancestors to all life on this planet (Kutschera and Niklas 2004).<\/p>\n<h2 class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">What Darwin was Missing<\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.527999877929688pt;margin-right: 2.886962890625pt;text-indent: 0pt\">Although the theory of evolution by natural selection gained traction in scientific circles in the decades after Darwin\u2019s publication of <em>Origins<\/em>, he was never able to discover the mechanisms that caused variation within members of the same species or the means by which traits were inherited. This began later in 1892 with the publication of <em>The Germ-Plasm: A Theory of Heredity<\/em> by August Weismann, the same Weismann of the mouse tail experiment presented earlier in this chapter. In his book, Weissman proposed a theory of germ-plasm, which was a precursor to the later discovery and understanding of DNA. Weismann specialized in cytology, a branch of biology devoted to understanding the function of plant and animal cells. Germ-plasm, he proposed, was a substance in the germ cells (what we would call gametes, or sex cells, today) that carried hereditary information. He predicted that an offspring inherits half of its germ-plasm from each of its parents, and claimed that other cells (e.g. somatic, or body, cells) could not transmit genetic information from parents to offspring. This thereby erased the possibility that acquired traits (which he argued resided in somatic cells) could be inherited (Zou 2015). This contribution to evolutionary theory was an important step toward understanding genetic inheritance, but a connection between genetics and evolution was still lacking.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 2.886962890625pt;text-indent: 0pt\"><span style=\"background-color: #ff9900\">A series of lectures by a deceased Augustinian monk named Gregor Mendel (1822\u20131884), originally published in 1865, changed that perspective (Moore 1993, 285). Although Darwin was unknown to Mendel, he began a series of experiments with pea plants shortly after the publication of Darwin\u2019s <em>Origins<\/em>, aiming to add to evolutionary understandings of heredity. As Mendel bred different generations of pea plants that had differences in seed shape and color, pod shape and color, flower position, and stem length, he documented consistent expression of some variations over others in subsequent generations. He meticulously documented the statistics of each crossing of plants and the percentages of <strong>phenotypes<\/strong> that resulted, eventually discovering the concept of dominance and recessiveness of characteristics. He also documented that there is no blending of inherited characteristics. For example, pea pod colors in the offspring of two parent plants, one with yellow pods and one with green, were <em>either<\/em> yellow or green, not yellowish green. Mendel also discovered that characteristics are inherited and expressed independently of each other, meaning the color of the pea pod was not necessarily expressed in conjunction with the pod being wrinkled or smooth. The recognition of the importance of Mendel\u2019s work began with its rediscovery by Hugo de Vries and Carl Correns, both of whom were working on hypotheses regarding heredity in plants and had arrived at conclusions similar to Mendel\u2019s. Both published papers supporting Mendel\u2019s conclusions in 1900 (Moore 1993). Research into the inheritance of characteristics continued through the next three decades, and by the close of the 1930s, no major scientific questions remained regarding the transmission of heredity through <strong>genes<\/strong>, although what genes did and what chemicals they were made of were still under investigation.\u00a0 <\/span><span style=\"text-decoration: underline\">(chapter 3)<\/span><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 2.886962890625pt;text-indent: 0pt\">The <strong>Modern Synthesis <\/strong>refers to the merging of Mendelian genetics with Darwinian evolution that took place between 1930 and 1950. The basic principles of the synthetic theory were influenced by scientists working in many different fields, including genetics, zoology, biology, paleontology, botany, and statistics. Although there were differences of opinion among them, evolution came to be defined as changes in allele frequencies within populations. Genetic mutations, changes in the genetic code that are the original source of variation in every living thing, were believed to be random, the sources of phenotypic variation, and transmitted through sexual reproduction. These assertions were supported by a growing body of field and laboratory research, as well as new work in mathematics in the field of population genetics that defined evolution as numerical changes in gene frequencies within an interbreeding population from one generation to the next (Corning 2020). These changes in gene frequencies were argued to be the result of natural selection, mutation, migration (<strong>gene flow<\/strong>), and <strong>genetic drift<\/strong>, or random chance. Empirical research and mathematics demonstrated that very small selective forces acting over a relatively long time were able to generate substantial evolutionary change, including speciation (Plutynski 2009). Thus, the Modern Synthesis encompassed both <strong>microevolution<\/strong>, which refers to changes in gene frequencies between generations within a population, and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1507\">macroevolution<\/a><\/strong>, longer-term changes in a population that can eventually result in speciation, wherein individuals from different populations are no longer able to successfully interbreed and produce viable offspring.<\/p>\n<figure style=\"width: 267px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image10-2.jpg\" alt=\"A white man with short hair dressed in a white shirt and dark tie.\" width=\"267\" height=\"355\" \/><figcaption class=\"wp-caption-text\"><em>Figure 2.13: Theodosius Dobzhansky (1943). Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Dobzhansky_no_Brasil_em_1943.jpg\">Dobzhansky no Brasil em 1943<\/a> by unknown creator via <a href=\"https:\/\/www.flickr.com\/photos\/celycarmo\/\">Cely Carmo<\/a> at Flickr is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/em><\/figcaption><\/figure>\n<p><em>Genetics and the Origin of Species<\/em>, published in 1937 by Theodosius Dobzhansky (Figure 2.13), was a cornerstone of the modern synthesis, applying genetics to the study of natural selection in wild populations, appealing to both geneticists and field biologists. Dobzhansky was interested in <strong>speciation<\/strong>, particularly in finding out what kept one species distinct from another and how speciation occurred. His research involved fruit flies, the species <em>Drosophila pseudoobscura<\/em>. At the time he began in the 1920s, biologists assumed all members of the same species had nearly identical genes. Dobzhansky traveled from Canada to Mexico capturing wild members of <em>D.<\/em><em>pseudoobscura<\/em>, discovering that different populations had different combinations of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_738\">alleles<\/a><\/strong> (forms of a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_382_1458\">gene<\/a><\/strong>) that distinguished them from other populations, even though they were all members of the same species. What, then, led to the creation of new species? Dobzhansky realized it was sexual selection. Members of the same species are more likely to live among their own kind and to recognize, and prefer, them as mates. Over time, as a result of random mutations, natural selection in a given environment, and <strong>genetic drift<\/strong>, meaning random changes in allele frequencies, members of the same population accumulate mutations distinct to their own <strong>gene pool<\/strong>, eventually becoming genetically distinct from other populations. What this means is that they have become a new <strong>species.<\/strong><\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.527999877929688pt;margin-right: 2.886962890625pt;text-indent: 0pt\">From these studies, Dobzhansky and others developed the Bateson-Dobzhansky-Muller model, also known as Dobzhansky-Muller model (Figure 2.14). It is a model of the evolution of genetic incompatibility. Combining genetics with natural selection, the model is important in understanding the role of reproductive isolation during speciation and the role of natural selection in bringing it about. Due to sexual selection (mate preference), populations can become reproductively isolated from one another. Eventually, novel mutations may arise and be selected for in one or both populations, rendering members of each genetically incompatible with the other, resulting in two distinct species.<\/p>\n<figure style=\"width: 601px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image4-4.jpg\" alt=\"Dobzhansky-Muller Model producing hybrids with incompatible mutations. See caption for image details.\" width=\"601\" height=\"348\" \/><figcaption class=\"wp-caption-text\">Figure 2.14: The Dobzhansky-Muller Model: In the ancestral population the genotype is AABB. When two populations become isolated from each other, new mutations can arise. In one population uppercase A evolves into lowercase a, and in the other uppercase B evolves into lowercase b. When the two populations hybridize, it is the first time a and b interact with each other. When these alleles are incompatible, they represent Dobzhansky-Muller incompatibilities. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Bateson-Dobzhansky-Muller_model._.jpg\">Bateson-Dobzhansky-Muller model<\/a> by OrientationEB is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Special Topic: Evolution and Natural Selection Observable Today<\/h2>\n<figure style=\"width: 339px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image19-3.jpg\" alt=\"Side view of a brown speckled lizard laying on a plastic lawn chair.\" width=\"339\" height=\"226\" \/><figcaption class=\"wp-caption-text\">Figure 2.15: Puerto Rican Crested Anole photographed in Picard, Dominica. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Anolis_cristatellus_in_Picard,_Dominica-2012_02_15_0339.jpg\">Anolis cristatellus in Picard, Dominica-2012 02 15 0339<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Postdlf\">Postdif<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/deed.en\">CC BY-SA 3.0 Unported License<\/a>.<\/figcaption><\/figure>\n<p>Although this chapter primarily focuses on the past, it is important to remember that natural selection and evolution are still ongoing processes. Climate change, deforestation, urbanization, and other human impacts on the planet are influencing evolution among many contemporary species of plants and animals. One such example occurs among crested anoles (<em>Anolis cristatellus<\/em>), small lizards of the Caribbean jungle that are increasingly making their home in cities (Figure 2.15).<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">As urban sprawl continues across the planet, shrinking the availability of wilderness habitat, many wild species have come to make their homes in cities. \u201cUrbanization has dramatically transformed landscapes around the world\u2014changing how animals interact with nature, creating \"heat islands\" with higher temperatures, and hurting local biodiversity. Yet many organisms survive and even thrive in these urban environments, taking advantage of new habitats created by humans (National Science Foundation 2023). A recent example of lizards in Puerto Rico demonstrates evolution happening quickly in both behavior and genes that has come about as a result of the pressures of urban life (Winchell et al. 2023). Crested anoles, who once lived only in forests, now scurry around towns and cities throughout the Caribbean. As a result of having to sprint across large open spaces, like hot streets and parking lots, they have developed longer limbs. City-living lizards also now sport longer toe pads with special scales that allow them to cling to smooth surfaces, like windows and walls (as well as the plastic patio furniture pictured in Figure 2.15), rather than to the rough surfaces of bark and rock that their forest-living relatives climb. These adaptations enhance their ability to escape predators and survive in cities.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Researchers were curious to see if these changes were the result of genetic changes in urban populations, so they captured 96 male lizards in three Puerto Rican regions and compared their genomes to each other and to forest specimens in each location. They found that members of the three city-living populations were genetically distinct from each other, as well as from forest populations in their respective regions. In total, 33 genes in the urban lizards\u2019 genomes were different from their forest-living counterparts and were linked to urbanization. These changes are estimated to have occurred just within the last 30 to 80 generations, suggesting that selective pressures related to survival in urban environments is strong. As study coauthor Kristen Winchell put it, \u201cWe are watching evolution as it is unfolding\u201d (National Public Radio 2023). (If you are interested in hearing more about the study, see \u201c<a href=\"https:\/\/www.pnas.org\/post\/podcast\/lizards-adapt-urban-living\">How Lizards Adapt to Urban Living<\/a>,\u201d an episode of Science Sessions, a free podcast from the Proceedings of the National Academy of Sciences (PNAS 2023) featuring Dr. Winchell and her work.)<\/p>\n<\/div>\n<h2 class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Misconceptions About Evolution Through Natural Selection<\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.836006164550781pt;margin-right: 3.1103515625pt;text-indent: 0pt\">After many years of teaching about evolution and natural selection, it continues to surprise me how many misconceptions exist about how the process works. If you do a web image search for \u201chuman evolution,\u201d the following image is likely to appear (Figure 2.16).<\/p>\n<figure style=\"width: 605px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image2-6.png\" alt=\"Six increasingly upright figures walk in one direction.\" width=\"605\" height=\"222\" \/><figcaption class=\"wp-caption-text\">Figure 2.16: An artist\u2019s visual representation of the process of human evolution. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Human_evolution_scheme.svg\">Human evolution scheme<\/a> by M. Garde is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 3.0211181640625pt;text-indent: 0pt\">What is wrong with this picture? First, it implies that humans evolved from chimpanzees, which is incorrect. Although, as primates, we share a common ancestor very far back in time, we split from other primates, including our closest relatives, the nonhuman apes, several million years ago. This image also suggests that evolution is gradual and progressive; that it is intentional and directional; and that there is an end to it\u2014a stopping point. As you will be learning, evolution takes place in fits and starts, depending on the physical environment, changes in climate, food supply, predation, reproductive success, and other factors. It is also not intentional, in the sense that there is no predetermined end; in fact, if environmental conditions change, species can evolve in different directions or even go extinct. Evolution also does not necessarily progress in the same direction over time. One example is the eel-like creature <em>Qikiqtania wakei<\/em> that lived 375 million years ago. It was originally a fish that evolved to walk on land, then evolved to live back in the water. Early tetrapods like <em>Qikiqtania<\/em> were likely spending more and more time out of the water during this period. The arrangement of bones and joints in their fins was starting to resemble arms and legs, which would have allowed them to prop themselves up in shallow water and survive on mudflats. <em>Qikiqtania\u2019<\/em>s skeletal morphology, however, suggests that it then evolved from having rudimentary fingers and toes back to fins that allowed them to again swim in open water (Stewart et al. 2022).<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.869003295898438pt;margin-right: 11.6299438476562pt;text-indent: 0pt\">There is also the misperception that natural selection can create entirely new anatomical structures out of thin air in response to changes in environmental pressures. For example, when asked if they can think of ways in which modern humans are continuing to evolve biologically, students often postulate that, as a result of climate change, humans might rapidly develop gills, webbed hands and feet, and learn to breathe underwater in response to rising sea levels. Unfortunately, natural selection can only act on slight variations in anatomy that are already present, and we have no rudimentary physiological system for breathing underwater. Given that natural selection can only act upon existing variation, humans have evolved in such a way that many parts of our bodies are prone to injury. Our knees are one example. The anterior cruciate ligament (ACL) in our knees is \u201cvulnerable to tearing in humans because our upright bipedal posture forces it to endure much more strain than it is designed to\u201d (Lents 2018, 23). When our ancestors made the transition from quadrupedalism to upright walking, we shifted from four bent legs to two straight legs, relying more on our bones than our muscles to support our weight. This is functional for normal walking and running in a straight line, but sudden shifts in direction and momentum, combined with the sizes and weights of modern humans, result in tears in an ACL that is simply not strong enough to bear the stress. If evolution had the capability to engineer a knee from scratch, it would look quite different, and any ligaments involved would likely be larger, stronger, and more flexible. For an interesting look at what anatomically modern humans might look like if we had evolved to withstand the stresses our bodies undergo in our present environment, see \u201c<a href=\"https:\/\/www.radiotimes.com\/tv\/documentaries\/this-is-what-the-perfect-body-looks-like-according-to-science\/\">This is what the perfect body looks like - according to science<\/a>,\u201d which was proposed by biological anthropologist Alice Roberts (Harrison 2018).<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.836006164550781pt;margin-right: 9.61920166015625pt;text-indent: 0pt\">Another misperception about evolution is that some species are \u201cmore evolved\u201d than others. Every species currently alive on the planet today is the result of millennia of natural selection that has rendered current members of that species well-adapted to their respective environments. Humans are no more \u201cevolved\u201d than fruit flies or yeast. What sets us apart are our cultural and technological abilities, which have allowed us to successfully survive in a wide variety of physical environments, many of which are now becoming too hot, too wet, or too dry to sustain human life without a great deal of technological intervention (IPCC 2022).<\/p>\n<figure style=\"width: 366px\" class=\"wp-caption alignright\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image5-3.jpg\" alt=\"A male dragonfly with a light blue body, transparent wings, and black markings rests on a twig.\" width=\"366\" height=\"282\" \/><figcaption class=\"wp-caption-text\">Figure 2.17: Adult male Common Whitetail Dragonfly, <a href=\"https:\/\/commons.wikimedia.org\/wiki\/Libellula_lydia\">Libellula lydia<\/a>. Credit: <a href=\"https:\/\/www.cirrusimage.com\/dragonfly_common_whitetail.htm\">Common Whitetail Dragonfly \u2013 Plathemis lydia<\/a> by Bruce Marlin is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.5\">CC BY-SA 2.5 License<\/a>.<\/figcaption><\/figure>\n<p>There is also some confusion about what \u201cfitness\u201d actually means and a failure to grasp that it changes as environmental conditions change. Evolutionary \u201cfitness\u201d is different from physical fitness. \u201cFitness\u201d in evolutionary terms refers to an individual\u2019s ability to survive and reproduce viable offspring who also survive and reproduce. Evolutionary fitness and reproductive success are highly dependent on specific environmental conditions, which can shift over time, greatly affecting the relative fitness of individuals in a population. Recent research on the impacts of climate change on dragonflies will serve to illustrate the point (Figure 2.17).<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.748001098632812pt;margin-right: 3.8743896484375pt;text-indent: 0.594001770019531pt\">Pictured here is a male dragonfly, who, you will notice, has distinctive black markings on its wings. This is due to melanization. Males control breeding, and those with more ornamentation tend to attract more mates and to successfully ward off male competitors. Higher levels of melanization, however, have negative consequences for males in warming climates. The black markings absorb heat, elevating body temperatures, which can cause overheating, reduce male fighting ability, and even lead to death (Moore et al. 2021). Females are not as adversely affected because they spend more time in shaded areas, while males are more often flying in sunlit areas, fending off rivals. However, as highly melanized males become less viable, wing coloration is undergoing selection in males. In other words, what constitutes being \u201cfit\u201d for males has changed, favoring those who have fewer of the black markings and, therefore, are less negatively impacted by warming temperatures. Note that natural selection acts on individuals, \u201cselecting\u201d those who happen to be fit for particular environmental conditions at a particular point in time. Evolution, though, happens at the level of the population. If the climate continues to warm, populations of dragonflies who inhabit warming areas will increasingly exhibit less ornamentation in males.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.858001708984375pt;margin-right: 5.50946044921875pt;text-indent: 0pt\">Lastly, natural selection can only act on characteristics that influence reproductive success. Deleterious traits that have nothing to do with one\u2019s ability to reproduce and successfully rear offspring to reproductive age will continue to be passed on. For example, the author of this chapter is a natural redhead, and redheads are predisposed genetically to a number of conditions that can negatively affect health (Colliss Harvey 2015), but some of these conditions are not diagnosed until later in life. One example is Parkinson\u2019s disease (Chen et al. 2017), which is a degenerative neurological disorder. The average age of diagnosis of Parkinson\u2019s is 60 years of age, meaning redheads may encounter such a diagnosis well past childbearing age, having already passed on the genetic predisposition. Thus, Parkinson\u2019s disease cannot be selected out from the redhead family tree.<\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.858001708984375pt;margin-right: 5.50946044921875pt;text-indent: 0pt\"><span style=\"background-color: #ff9900\">Dig Deeper: Teaching Evolution Around the World\u00a0 <\/span><span style=\"text-decoration: underline\">(Put in Chapter 17 or refer such a subject to chap 17)<\/span><\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-right: 0.885009765625pt\"><span style=\"background-color: #ff9900\">Evolution is recognized as a central organizing principle for all scientific disciplines and accepted without controversy among scientists and educated people around the world. The United States has historically been the exception (Lerner 2000). In some parts of the U.S. the teaching of evolution to K-12 students continues to evoke controversy, related to politics and religion. The problem is compounded by the degree of control individual states and local school boards exercise over curriculum in the nation\u2019s public schools (Lerner 2000).<\/span><\/p>\n<figure style=\"width: 515px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image21-1.jpg\" alt=\"A crowd of people face a fenced-off area outside a brick building. \" width=\"515\" height=\"361\" \/><figcaption class=\"wp-caption-text\"><span style=\"background-color: #ff9900\">Figure 2.18: Crowds attend the Scopes trial, Dayton, Tennessee, July 20, 1925. Credit: <a style=\"background-color: #ff9900\" href=\"https:\/\/siarchives.si.edu\/collections\/siris_arc_308484\">Clarence S. Darrow interrogating William Jennings Bryan, Scopes trial<\/a> (1925) by William Silverman via <a style=\"background-color: #ff9900\" href=\"https:\/\/www.flickr.com\/photos\/smithsonian\/\">Smithsonian Institution<\/a> has <a style=\"background-color: #ff9900\" href=\"https:\/\/www.si.edu\/termsofuse\">no known copyright restrictions.<\/a> [Smithsonian Institution archives, Acc. 10-042, William Silverman Photographs, 1925, Image ID: 2009-21077.]<\/span><\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;margin-right: 0.6488037109375pt;text-indent: 0pt\"><span style=\"background-color: #ff9900\">The debate over teaching evolution in schools in the United States first came to a head in 1925 after several states attempted to legislate a ban against its teaching. The state of Tennessee eventually did pass such a ban, and the American Civil Liberties Union (ACLU) offered to defend any science teacher who agreed to break the law. John Scopes, who taught in a small, rural Tennessee school, continued to teach evolution, resulting in the \u201cScopes Monkey Trial,\u201d one of the most famous media trials in American history (Figure 2.18). The entire nation listened to its broadcast live on the radio and read about it daily in hundreds of newspapers. Scopes was defended by Clarence Darrow, the most famous lawyer in the country at the time; Scopes was eventually convicted, though the conviction was later overturned. The pro-Evolution movement benefited greatly from Darrow\u2019s questioning of those on the anti-Evolution side, whose responses were perceived negatively by well-educated listeners in northern cities. The teaching of evolution was also bolstered by a Supreme Court decision in 1947 overriding states\u2019 rights to make decisions on church-state issues and by the launching of Sputnik, the first Soviet (Russian) satellite, in 1957, making science education a national priority. During the 1960s to 1970s, <strong>creationism<\/strong> and <strong>intelligent design<\/strong> began to take hold as courts ruled in favor of \u201cacademic freedom\u201d (Pew Research Center 2019). Since that time, states, and even local school boards, have pushed for the removal of evolution from science curriculum and textbooks or for teaching evolution on equal footing with concepts such as creationism and intelligent design (Masci 2019).<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ff9900\">Partly in response to chronically low scores by students in the United States on international measures of ability in science, math, and reading (Desilver 2017), development of the Next Generation Science Standards (NGSS) for K-12 education began in 2011 and were implemented in 2013 in public schools across the nation (nextgenscience.org). The standards were developed collaboratively by The National Research Council (NRC; a branch of the National Academy of Sciences), the National Science Teachers Association, the American Association for the Advancement of Science, and Achieve, an independent, nonpartisan, nonprofit education-reform organization dedicated to working with states to raise academic standards and graduation rates. Twenty-six states also partnered together in the development of the standards. However, due to a number of issues, including funding and support for teachers, adoption of the standards has varied by state. States were also allowed to alter the curriculum. One of the main issues in several states with the original curriculum was the teaching of evolution (Pew Research Center 2014). One prominent aspect to note is that the NGSS do not specify that human evolution be taught, and high school standards do not require teaching about all of the forces of evolution, only mutation and natural selection.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ff9900\">The situation regarding teaching evolution has changed greatly in recent years. A 2007 survey found that only one in three public high school biology teachers in the United States presented evolution consistently with the recommendations of the nation\u2019s leading scientific authorities, and 13% of these teachers emphasized creationism as a valid scientific alternative to modern evolutionary biology (Plutzer, Branch, and Reid 2020). A repeat of the survey in 2019 demonstrated marked improvement in the amount of time teachers devoted to teaching evolution, as well as more teacher training and preparedness to teach evolution. Such improvements were attributed to the need to meet the Next Generation Science Standards, as well as continuing outreach by the National Science Teaching Association, the National Association of Biology Teachers, and the National Academy of Sciences in producing classroom resources and providing professional development opportunities to advance the inclusion of evolution in the nation\u2019s classrooms.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ff9900\">Public acceptance of evolution has also substantially improved in recent years (Miller et al. 2022). National samples of American adults have been asked at regular intervals since 1985 to agree or disagree with the following statement: \u201cHuman beings, as we know them today, developed from earlier species of animals (Miller et al. 2022)\u201d During the last decade, the percentage of U.S. adults agreeing with this statement increased from 40% to 54%\u2014a majority for the first time. This level of acceptance of evolution in the United States is atypically low for a developed nation. In a study of the acceptance of evolution in 34 developed nations in 2005, only Turkey\u2014at 27%\u2014scored lower than the United States (Miller, Scott and Okamoto 2006). There are also distinct differences among members of the U.S. population in terms of acceptance of evolution, with 68% of those ages 18\u201324, 58% of those with college degrees, and 65% of those who have taken four or more college-level science courses the most accepting of evolution. An increasing number of parents also report changing their unfavorable views of evolution due to helping their children with science homework and science fair projects.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ff9900\">Approaches to teaching evolution vary across the globe, with considerable differences within and between nations. One example is the United Kingdom, home of Darwin and Wallace. There, evolution is not introduced until ages 14\u201316, which is considered quite late by some educators. And, although evolution is taught in biology classes, it is addressed as a separate topic, rather than integrated into the curriculum as a foundational concept. As in the United States, there is also considerable variation between public and private schools, and between religious and secular institutions, in their treatment of the topic and the inclusion of alternative viewpoints, such as creationism (Harmon 2011). There are similar differences across the European Union, including within different populations in member countries.<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ff9900\">There are also differences in the teaching of evolution across many predominantly Muslim nations. Salman Hameed, Professor of Integrated Science and Humanities at Hampshire College and Director of the Center for the Study of Science in Muslim Societies (SSiMS), whose research focuses on the acceptance of evolution among Muslims, has uncovered a great deal of variation among Muslims regarding beliefs about evolution. He points out that there is no central position within Islam regarding evolution, leaving it up to governments, textbook authors, and other entities to decide whether, and how, to address evolution in education. Saudi Arabia, Oman, Algeria, Morocco, and Lebanon all ban the teaching of evolution on religious grounds. Other Islamic nations, including Egypt, Malaysia, Syria, and Turkey, include evolutionary concepts like natural selection in their science curriculum but refrain from discussing human evolution (Asghar, Hameed, and Farahani 2014).<\/span><\/p>\n<p class=\"import-Normal\"><span style=\"background-color: #ff9900\">Impeding acceptance of evolution in science classes around the world is the adoption of textbooks from the E.U., the U.K., and the U.S. that include examples that are not culturally relevant to local populations (Harmon 2011). One classic example Hameed cites is that of the peppered moths in England whose predominant color pattern evolved from mostly white to mostly black due to pollutants darkening the tree bark of their habitat during the Industrial Revolution. Historical examples like this have little relevance for 21st-century students who grew up in non-Western countries and know little of England\u2019s history or of the species that live there. It also privileges Western science over local science, to which many individuals in former European colonies and territories object (Jones 2017). Hameed suggests customizing textbooks to include local fossils and species whenever possible (Harmon 2011).<\/span><\/p>\n<\/div>\n<\/div>\n<h2 class=\"__UNKNOWN__\">Are We Still Evolving?<\/h2>\n<div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">After reading this chapter, many students are curious to know if humans are still evolving. The answer is yes. As a species, we continue to respond to selective pressures biologically and culturally. This final section will focus on three contemporary examples of human evolution. Before beginning, let\u2019s review the conditions necessary for natural selection to operate on a trait. First, the trait must be heritable, meaning it is transmitted genetically from generation to generation. There must also be variation of the trait within the population and the trait must influence reproductive success. Three examples of traits that meet these criteria are immunity to the Human Immunodeficiency Virus (HIV), height, and wisdom teeth (Andrews, Kalinowski, and Leonard 2011).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">AIDS is a potentially fatal infectious disease caused by HIV, a zoonosis believed to be derived from Simian Immunodeficiency Viruses (SIVs) found in chimpanzees and monkeys and most likely transmitted to humans through the butchering of infected animals (Sharp and Hahn 2011). In total, 40 million people have died from AIDS-related illnesses since the start of the global epidemic in the 1980s. There were 38.4 million people around the world living with AIDS as of 2021, including 1.5 million new cases and 650,000 deaths in that year alone (UNAIDS 2021). A disease causing this level of morbidity and mortality represents a major selective pressure, especially given that infection can occur before birth (Goulder et al. 2016), thereby affecting future reproductive success.<\/p>\n<figure style=\"width: 323px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image17-7.png\" alt=\"A political map of Europe and North Africa associated with percentages ranging from 0% to 16.4%.\" width=\"323\" height=\"391\" \/><figcaption class=\"wp-caption-text\">Figure 2.19: Map of CCR5-delta32 allele distribution. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-11\/\">Map of CCR5-delta32 allele distribution (Figure 16.10)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Katie Nelson is a collective work under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. <a href=\"https:\/\/docs.google.com\/document\/d\/1VUDKMBJYS_jNONjLxT04jQN0_z9Ua50BRN6auGSHUuU\/edit\" target=\"_blank\" rel=\"noopener\">A full text description of this image is available.<\/a> [Includes <a href=\"https:\/\/pixabay.com\/vectors\/europe-map-western-political-32847\/\">Europe Map Western Political 32847<\/a> by <a href=\"https:\/\/pixabay.com\/users\/clker-free-vector-images-3736\/\">Clker-Free-Vector-Images<\/a>, <a href=\"https:\/\/pixabay.com\/service\/terms\/#license\">Pixabay License<\/a>; data from Solloch et al. 2017.]<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The majority of people in the world are highly susceptible to HIV infection, but some are not. These latter individuals are homozygous for a rare, recessive allele at the CCR5 locus that makes them immune to HIV. Heterozygotes who inherit a single copy of this allele are more resistant to infection and, when infected, the disease takes longer to progress in the event that they are infected. The mechanism by which the allele prevents infection involves a 32-base pair deletion in the DNA sequence of the CCR5 gene, creating a nonfunctioning receptor on the surface of the cell that prevents HIV from infecting the cell. The allele is inherited as a simple Mendelian trait, and there is variation in its prevalence, ranging as high as 14% of the population in northern Europe and Russia (Novembre, Galvani, and Slatkin 2005; see Figure 2.19). What is interesting about the allele\u2019s geographic distribution is that it does not map onto parts of the world with the highest rates of HIV infection (Figure 2.20), suggesting that AIDS was not the original selective pressure favoring this allele (see Figures 2.19 and 2.20).<\/p>\n<figure style=\"width: 498px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image7-8.png\" alt=\"World map with different HIV infection rates throughout the world.\" width=\"498\" height=\"253\" \/><figcaption class=\"wp-caption-text\">Figure 2.20: World map of countries shaded according to their HIV\/AIDS adult prevalence rate in 2020. <a href=\"https:\/\/docs.google.com\/document\/d\/1VUDKMBJYS_jNONjLxT04jQN0_z9Ua50BRN6auGSHUuU\/edit\" target=\"_blank\" rel=\"noopener\">A full text description of this image is available.<\/a> Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:World_map_of_countries_by_HIV-AIDS_adult_prevalence_rate_%282020%29.svg\">World map of countries by HIV-AIDS adult prevalence rate (2020)<\/a> by LuccaSSC has been designated to the <a href=\"https:\/\/creativecommons.org\/share-your-work\/public-domain\/cc0\/\">public domain (CC0 1.0)<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Given its current geographic distribution, the bubonic plague, which ravaged Europe repeatedly from the 14th to the 19th centuries (Pamuk 2007), was initially proposed as the selective agent. Subsequent research suggests smallpox, which killed up to 400,000 people annually in 18th-century Europe (Hays 2005), was more likely the selective pressure (Novembre, Galvani, and Slatkin 2005). Given the mortality rates for smallpox (Crosby 2003), an allele that conferred immunity was highly advantageous, as it is for those faced with the threat of HIV infection today.<\/p>\n<p class=\"import-Normal\">Height is another example of a trait experiencing selective pressure. If you have ever toured a historical site, you have likely hit your head on a doorframe or become claustrophobic trying to squeeze down a narrow hallway under a lower-than-average ceiling. It is not your imagination. Humans have gotten taller in recent centuries. In fact, the average height of people in industrialized nations has increased approximately 10 centimeters (about four inches) in the past 150 years. This increase has been attributed to improvements in nutrition, sanitation, and access to medical care (Hatton 2014). But this is only part of the story.<\/p>\n<p class=\"import-Normal\">Height is highly heritable. Studies indicate 80% of variation in height within populations is due to genetics, with 697 different genetic variances identified as having an effect on adult stature (Devuyst 2014). Multiple studies also demonstrate a positive relationship between height and reproductive success for men (Andrews, Kalinowsky, and Leonard 2011). This is primarily due to sexual selection and nonrandom mating, namely women\u2019s preferences for taller men, which may explain why height is one characteristic men often lie about on dating websites (Guadagno, Okdie, and Kruse 2012). Sexual selection also plays out with regard to economic success in Western cultures, with taller men more likely to be in higher-level positions that pay well. Research demonstrates an inch of height is worth an additional $789 per year in salary, meaning a man who is six feet tall will earn on average $5,525 more per year than an identical man who is five foot five purely due to heightism bias (Gladwell 2007). Over the course of a career, this can add up to hundreds of thousands of dollars, likely allowing taller men to attract more potential mates, increasing their reproductive success.<\/p>\n<p class=\"import-Normal\">Wisdom teeth are also undergoing evolutionary pressure. Have you or anyone in your family had their wisdom teeth removed? While it can be a painful and expensive process, it is a common experience in Western nations. It begs the question as to why there is no longer room in our mouths for all of our teeth? Biological anthropologist Daniel Lieberman offers several reasons, including that modern humans are growing faster and maturing earlier, which could be leading to impaction if skeletal growth takes place faster than dental growth. He also argues that the soft diets many modern humans consume generate insufficient strain to stimulate enough growth in our jaws to accommodate all of our teeth. Lastly, as the human brain has expanded over the past hundreds of thousands of years, it is taking up more space in the skull, causing the jaw to shrink, leaving no room for third molars (Lieberman 2011).<\/p>\n<p class=\"import-Normal\">Conversely, do you know anyone whose wisdom teeth never came in? That is fairly common in some populations, suggesting evolutionary pressure favoring the absence of wisdom teeth has been culturally influenced. The oldest fossil evidence of skulls missing third molars was found in China and is 300,000 to 400,000 years old, suggesting the earliest mutation selecting against the eruption of wisdom teeth arose in Asia (Main 2013). Since that time, jaws have continued to decrease in size to the point they often cannot accommodate third molars, which can become impacted, painful, and even infected, a condition physical anthropologist Alan Main argues might have interfered with the ability to survive and reproduce in ancestral populations (Main 2013). As we have learned, a mutation that positively influences reproductive success\u2014such as being born without the trait to develop wisdom teeth\u2014would likely be selected for over time. Evidence in modern humans suggests that this is the case, with 40% of modern Asians and 45% of Native Alaskans and Greenlanders (populations descended from Asian populations) lacking wisdom teeth. The percentage among those of European descent ranges from 10 to 25% and for African Americans is 11% (Main 2013). Later chapters of this textbook emphasize that directional selection progresses along a particular path until the environment changes and a trait is no longer advantageous. In the case of wisdom teeth, the ability of modern dentistry to preempt impaction through surgery may, in fact, be what is preventing natural selection from doing away with wisdom teeth altogether.<\/p>\n<h2 class=\"import-Normal\" style=\"margin-left: 1.364006042480469pt;text-indent: 0pt\">Key Developments in Evolutionary Thought<\/h2>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 482.25pt\">\n<tbody>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">4th century BCE<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Aristotle<\/p>\n<p class=\"import-Normal\">(384\u2013322 BCE)<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">\u201cFounder of Biology.\u201d Publishes <em>History of Animals<\/em>, a biological classification system of over 500 animals based on structure, physiology, reproduction, and behavior. Also creates the \u201cGreat Chain of Being,\u201d ranking species and placing humans closest to God.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">8th\u20139th century CE<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Al-Jahiz<\/p>\n<p class=\"import-Normal\">(776\u2013868 CE)<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Writes seven-volume <em>Book of Animals<\/em>, which includes animal classifications and food chains. Introduces concept of biological evolution and its mechanisms.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">1011\u20131021<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Ibn al-Haythem<\/p>\n<p class=\"import-Normal\">(965\u20131040 CE)<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">\u201cFather of Modern Optics.\u201d Uses experimental science to catalog how vision works and discovers laws of reflection and refraction. Publishes <em>Book of Optics<\/em> and invents <em>camera obscura<\/em>, the foundation for modern photography.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">1620<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Francis Bacon<\/p>\n<p class=\"import-Normal\">(1561\u20131626)<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">\u201cFather of Empiricism.\u201d Publishes <em>The Novum Annum<\/em>, formulating the scientific method based on observation and inductive reasoning.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">1686<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">John Ray<\/p>\n<p class=\"import-Normal\">(1627\u20131705)<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">First to publish a biological definition of <em>species<\/em> in <em>History of Plants<\/em>.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">1749<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Comte de Buffon<\/p>\n<p class=\"import-Normal\">(1707\u20131788)<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Publishes <em>Histoire Naturelle<\/em>, comparing anatomical structures across species using methods still in use today. Inspires Lamarck and Cuvier.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">1758<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Carl von Linne<\/p>\n<p class=\"import-Normal\">(Carolus Linnaeus)<\/p>\n<p class=\"import-Normal\">(1707\u20131778)<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Introduces system of binomial nomenclature. Publishes <em>Systema Naturae<\/em>, the tenth edition of which introduces the designation <em>Homo sapiens <\/em>for humans.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">1788<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">James Hutton<\/p>\n<p class=\"import-Normal\">(1726\u20131797)<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">\u201cFather of Geology.\u201d Publishes <em>Theory of the Earth<\/em>; introduces idea of Deep Time; explains how features of the earth were formed through the actions of rain, wind, rivers, and volcanic eruptions.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">1798<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Thomas Malthus<\/p>\n<p class=\"import-Normal\">(1766\u20131834)<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Economist and \u201cFather of Statistics.\u201d Publishes <em>An Essay on Population<\/em>; introduces concept of carrying capacity; explains how populations outstrip the food supply, leaving some individuals to die off; inspires Darwin\u2019s idea of \u201cnatural selection.\u201d<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">1809<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Jean-Baptiste Lamarck<\/p>\n<p class=\"import-Normal\">(1744\u20131829)<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Publishes theory of the Inheritance of acquired characteristics; is the first Western scientist to propose a mechanism explaining how traits change in species over time and to recognize the importance of the physical environment in acting on species and their survival.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">1810<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Georges Cuvier<\/p>\n<p class=\"import-Normal\">(1769\u20131832)<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Paleontologist\/comparative anatomist; proved species went extinct; proposed the Theory of Catastrophism.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">1830<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Charles Lyell<\/p>\n<p class=\"import-Normal\">(1797\u20131875)<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Establishes geology as a science. Publishes first edition of <em>The Principles of Geology <\/em>(1830\u201333); issuing 12 total editions in his lifetime, each updated according to new scientific data.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">1858<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Alfred Russel Wallace<\/p>\n<p class=\"import-Normal\">(1823\u20131913)<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Sends scientific paper to Darwin titled \u201cOn the Tendency of Varieties to Depart Indefinitely from the Original Type,\u201d essentially espousing the concept of natural selection; a reading of the papers by both Wallace and Darwin to the Linnaean Society is conducted by Lyell.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">1859<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Charles Darwin<\/p>\n<p class=\"import-Normal\">(1809\u20131882)<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Publishes <em>On the Origin of Species by Means of Natural Selection<\/em> (1859).<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">1865<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Gregor Mendel<\/p>\n<p class=\"import-Normal\">(1822\u20131884)<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Publishes <em>Experiments in Plant Hybridization<\/em> (1865), outlining the fundamentals of genetic inheritance.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">1889<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">August Weismann<\/p>\n<p class=\"import-Normal\">(1834\u20131914)<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Publishes <em>Essays Upon Heredity <\/em>(1889), disproving the inheritance of acquired characteristics. Publishes <em>The Germ Plasm <\/em>(1892), postulating an early idea of inheritance through sexual reproduction.<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 0\">\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">1937<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Theodosius Dobzhansky<\/p>\n<p class=\"import-Normal\">(1900\u20131975)<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">One of the founders of the Modern Synthesis of biology and genetics. Publishes <em>Genetics and the Origin of Species<\/em> (1937). Documents a genetic model of speciation through reproductive isolation.<\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Review Questions<\/h2>\n<ul>\n<li class=\"import-Normal\">Summarize the major scientific developments that led to the formulation of the theory of natural selection.<\/li>\n<li class=\"import-Normal\">Explain how natural selection operates and how it leads to evolution in populations.<\/li>\n<li class=\"import-Normal\">Explain the importance of genetics to an understanding of human evolution.<\/li>\n<li class=\"import-Normal\">Have you observed current examples of evolution taking place where you live? In which species? Which forces of evolution are involved?<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<h2 class=\"__UNKNOWN__\">Key Terms<\/h2>\n<div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.902000427246094pt;margin-right: 18.1800537109375pt;text-indent: 0.406997680664062pt\"><strong>Allele<\/strong>: A nonidentical DNA sequence found in the same gene location on a homologous chromosome, or gene copy, that codes for the same trait but produces a different phenotype.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.836006164550781pt;margin-right: 4.92626953125pt;text-indent: 0.34100341796875pt\"><strong>Artificial selection<\/strong>: The identification by humans of desirable traits in plants and animals, and the subsequent steps taken to enhance and perpetuate those traits in future generations.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.836006164550781pt;margin-right: 0.62152099609375pt;text-indent: 0.461997985839844pt\"><strong>Binomial nomenclature<\/strong>: A system of classification in which a species of animal or plant receives a name consisting of two terms: the first identifies the genus to which it belongs, and the second identifies the species.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 6.76727294921875pt;text-indent: 0pt\"><strong>Carrying capacity<\/strong>: The number of living organisms, including animals, crops, and humans, that a geographic area can support without environmental degradation.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.924003601074219pt;margin-right: 3.10675048828125pt;text-indent: 0.0879974365234375pt\"><strong>Catastrophism<\/strong>: The theory that the Earth\u2019s geology has largely been shaped by sudden, short-lived, violent events, possibly worldwide in scope. Compare to uniformitarianism.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.924003601074219pt;margin-right: 3.10675048828125pt;text-indent: 0.0879974365234375pt\"><strong>Comparative anatomy<\/strong>: Georges-Louis Leclerc\u2019s technique of comparing similar anatomical structures across different species.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.924003601074219pt;margin-right: 3.10675048828125pt;text-indent: 0.0879974365234375pt\"><strong>Creationism<\/strong>: The belief that the universe and all living organisms originate from specific acts of divine creation, as in the Biblical account, rather than by natural processes such as evolution.<\/p>\n<p class=\"import-Normal\" style=\"margin-right: 0.0257568359375pt\"><strong>Empiricism<\/strong>: The idea that all learning and knowledge derives from experience and observation. It became prominent in the 17th and 18th centuries in western Europe due to the rise of experimental science.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.858001708984375pt;margin-right: 53.6096496582031pt;text-indent: 0.44000244140625pt\"><strong>Evolution<\/strong>: In a biological sense, this term refers to cumulative inherited change in a population of organisms through time. More specifically, <em>evolution<\/em> is defined as a change in allele (gene) frequencies from one generation to the next among members of an interbreeding population.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 1.298004150390625pt;margin-right: 0pt;text-indent: 0pt\"><strong>Extant<\/strong>: Still in existence; surviving.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 1.199005126953125pt;margin-right: 19.8264770507812pt;text-indent: 0.0989990234375pt\"><strong>Extinct<\/strong>: Said of a species, family, or other group of animals or plants that has no living members; no longer in existence.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 1.199005126953125pt;margin-right: 19.8264770507812pt;text-indent: 0.0989990234375pt\"><strong>Fixity of <\/strong><strong>Species<\/strong>: The idea that a species, once created, remains unchanged over time.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.836006164550781pt;margin-right: 6.576171875pt;text-indent: 0pt\"><strong>Gene<\/strong>: A sequence of DNA that provides coding information for the construction of proteins.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0.836006164550781pt;margin-right: 6.576171875pt;text-indent: 0pt\"><strong>Genetic drift<\/strong>: Random changes in allele frequencies within a population from one generation to the next.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.836006164550781pt;margin-right: 6.576171875pt;text-indent: 0pt\"><strong>Gene flow<\/strong>: The introduction of new genetic material into a population through interbreeding between two distinct populations.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0.836006164550781pt;margin-right: 6.576171875pt;text-indent: 0pt\"><strong>Gene pool<\/strong>: The entire collection of genetic material in a breeding community that can be passed from one generation to the next.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0.836006164550781pt;margin-right: 6.576171875pt;text-indent: 0pt\"><strong>Genotype<\/strong>: The genotype of an organism is its complete set of genetic material\u2014its unique sequence of DNA. Genotype also refers to the alleles or variants an individual carries in a particular gene or genetic location.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.693000793457031pt;margin-right: 1.15521240234375pt;text-indent: 0.605003356933594pt\"><strong>Hybrid<\/strong>: Offspring of parents that differ in genetically determined traits.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.924003601074219pt;margin-right: 16.4979858398438pt;text-indent: 0.319000244140625pt\"><strong>Intelligent design<\/strong>: A pseudoscientific set of beliefs based on the notion that life on earth is so complex that it cannot be explained by the scientific theory of evolution and therefore must have been designed by a supernatural entity.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.594001770019531pt;margin-right: 24.672607421875pt;text-indent: 0.681999206542969pt\"><strong>Macroevolution<\/strong>: Large and often-complex changes in biological populations, such as species formation.<\/p>\n<p><strong>Microevolution<\/strong>: Changes in the frequency of a gene or allele in an interbreeding population.<\/p>\n<p><strong>M<\/strong><strong>odern synthesis<\/strong>: The mid\u201320th century merging of Mendelian genetics with Darwinian evolution that resulted in a unified theory of evolution.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.693000793457031pt;margin-right: 3.12261962890625pt;text-indent: 0.583000183105469pt\"><strong>Natural selection<\/strong>: The natural process by which the survival and reproductive success of individuals or groups within an interbreeding population that are best adjusted to their environment leads to the perpetuation of genetic qualities best suited to that particular environment at that point in time.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.693000793457031pt;margin-right: 19.8097534179688pt;text-indent: 0.605003356933594pt\"><strong>Phenotype<\/strong>: The detectable or visible expression of an organism\u2019s <em>genotype<\/em>.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.693000793457031pt;margin-right: 7.392578125pt;text-indent: 0.198005676269531pt\"><strong>Scientific method<\/strong>: A method of procedure that has characterized natural science since the 17th century, consisting of systematic observation, measurement, experimentation, and the formulation, testing, and modification of hypotheses.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0.858001708984375pt;margin-right: 2.66143798828125pt;text-indent: 0.0330047607421875pt\"><strong>Speciation<\/strong>: The process by which new genetically distinct species evolve from the main population, usually through geographic isolation or other barriers to gene flow.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0.858001708984375pt;margin-right: 2.66143798828125pt;text-indent: 0.0330047607421875pt\"><strong>Species<\/strong>: A group of living organisms consisting of similar individuals capable of exchanging genes or interbreeding. The species is the principal natural taxonomic unit, ranking below a genus and denoted by a Latin binomial (e.g., <em>Homo sapiens<\/em>).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0.495002746582031pt;margin-right: 1.47552490234375pt;text-indent: 0pt\"><strong>Uniformitarian<\/strong><strong>ism<\/strong>: The theory that changes in the earth's crust during geologic history have resulted from the action of continuous and uniform processes\u2014such as wind, precipitation, evaporation, condensation, erosion, and volcanic action\u2014that continue to act in the present. Compare to <strong>c<\/strong><em>atastrophism<\/em>.<\/p>\n<h2 class=\"import-Normal\">About the Author<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 1.995002746582031pt;text-indent: 0pt\"><img class=\"alignleft\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image1-7.png\" alt=\"Person wearing outdoor gear including hat and sunglasses with mountains in the background.\" width=\"263\" height=\"351\" \/><\/p>\n<h3 class=\"import-Normal\">Joylin Namie, Ph.D.<\/h3>\n<p class=\"import-Normal\">Truckee Meadows Community College, <a class=\"rId106\" href=\"mailto:jnamie@tmcc.edu\">jnamie@tmcc.edu<\/a><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0.495002746582031pt;margin-right: 1.0625pt;text-indent: 0.847000122070312pt\">Joylin Namie is Professor of Anthropology at Truckee Meadows Community College, where she teaches courses in biological and cultural anthropology. Her current research interest is in (un)sustainable tourism in desert environments, particularly in the country of Jordan and the U.S. state of Nevada. She was awarded a fellowship to Jordan from the Council of American Overseas Research Centers (CAORC) in 2020 to explore this topic, including visiting Petra and other important tourism destinations in Jordan. Dr. Namie\u2019s favorite things in life are teaching, traveling, and spending time with her dog, Charley.<\/p>\n<p>&nbsp;<\/p>\n<p>&nbsp;<\/p>\n<h2 class=\"import-Normal\">For Further Exploration<strong><br \/>\n<\/strong><\/h2>\n<p>Costa, James T. 2017. <em>Darwin\u2019s Backyard: How Small Experiments Led to a Big Theory<\/em>. New York: W.W. Norton.<\/p>\n<p>Darwin, Charles. 1905. <em>The Voyage of the Beagle<\/em>. (Originally published in 1839 as <em>Journal and Remarks<\/em>). [Author\u2019s note: Several editions exist with different publishers, including illustrated editions, paperback editions, and e-books.]<\/p>\n<p>Moore, John A. 1993. <em>Science as a Way of Knowing: The Foundations of Modern Biology<\/em>. Cambridge, MA: Harvard University Press.<\/p>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 29.0806274414062pt;text-indent: 0pt\">Al-Haytham, Ibn. 1011-1021. <em>Kit\u0101b al-Man\u0101\u1e93ir<\/em> (Book of Optics). Cairo, Egypt.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 29.0806274414062pt;text-indent: 0pt\">Al-Jahiz. 776\u2013868 CE. <em>Kitab al-Hayawan<\/em> (<em>Book of <\/em><em>Animals<\/em><em>).<\/em><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Andrews, Tessa M., Steven T. Kalinowski, and Mary J. Leonard. 2011. \u201cAre Humans Evolving? A Classroom Discussion to Change Students\u2019 Misconceptions Regarding Natural Selection.\u201d <em>Evolution: Education and Outreach<\/em> 4 (3): 456\u2013466.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Aristotle. 384-322 BCE. <em>History of <\/em><em>Animals<\/em><em>.<\/em><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Asghar, Anila, Salman Hameed, and Najme Kashani Farahani. 2014. \u201cEvolution in Biology Textbooks: A Comparative Analysis of Five Muslim Countries.\u201d Religion &amp; Education 41 (1). Accessed February 12, 2023. https:\/\/www.tandfonline.com\/doi\/abs\/10.1080\/15507394.2014.855081.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Associated Press. January 10, 2023. \u201cForest Lizards Have Genetically Morphed To Survive Life In The City, Researchers Say.\u201d <em>National Public Radio (NPR)<\/em>. Retrieved February 19, 2023 from https:\/\/www.npr.org\/2023\/01\/10\/1148150056\/forest-lizards-genetically-morphed-cities.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 14.25pt;text-indent: 0pt\">Burkhardt, Richard W. 2013. \u201cLamarck, Evolution, and the Inheritance of Acquired Characters.\u201d <em>Genetics<\/em> 194 (4): 793\u2013805.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 21.4119873046875pt;text-indent: 0pt\">Chen, Xiqun, Danielle Feng, Michael A. Schwartzchild, and Xiang Gao. 2017. \u201cRed Hair, MC1R Variants, and Risk for Parkinson\u2019s Disease\u2014A Meta-Analysis.\u201d <em>Annals of Clinical and Translational Neurology<\/em> 4 (3): 212\u2013216.https:\/\/doi.org\/10.1002\/acn3.381.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 4.13275146484375pt;text-indent: 0pt\">Colliss Harvey, Jacky. 2015. <em>Red: A History of the Redhead<\/em>. New York: Black Dog &amp; Levanthal.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 4.13275146484375pt;text-indent: 0pt\">Corning, Peter A. 2020. \u201cBeyond the Modern Synthesis: A Framework for a More Inclusive Biological Synthesis.\u201d <em>Progress in Biophysics and Molecular Biology<\/em> 153: 5\u201312.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 2.188232421875pt;text-indent: 0pt\">Costa, James T. 2017. <em>Darwin\u2019s Backyard: How Small Experiments Led to a Big Theory<\/em>. New York: W. W. Norton.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 27.93212890625pt;text-indent: 0pt\">Crosby, Alfred W., Jr. 2003. <em>The Columbian Exchange: Biological and Cultural Consequences of 1492<\/em>. 30th Anniversary Edition. Westport, CT: Praeger.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 27.93212890625pt;text-indent: 0pt\">Darwin, Charles. 1859. <em>On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life. <\/em>First Edition. London: John Murray.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 35.9136962890625pt;text-indent: 0pt\">Darwin, Francis, ed. 2001[1897]. <em>The Life &amp; Letters of Charles Darwin<\/em>, vol. 2. Honolulu: University Press of the Pacific.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 7.79656982421875pt;text-indent: 0pt\">Desilver, Drew. 2017. \u201cU.S. Students\u2019 Academic Achievement Still Lags That of Their Peers in Many Other Countries.\u201d Pew Research Center, February 15. Accessed May 25, 2022. https:\/\/www.pewresearch.org\/fact-tank\/2017\/02\/15\/u-s-students-internationally-math-science\/.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Devuyst, Olivier. 2014. \u201cHigh Time for Human Height.\u201d <em>Peritoneal Dialysis International<\/em> 34 (7):685\u2013686.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Dobzhansky, Theodosius. 1937. <em>Genetics and the Origin of Species<\/em>. Columbia University Biological Series (Volume 11). New York: Columbia University Press.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 15.1101684570312pt;text-indent: 0pt\">Dunbar-Ortiz, Roxanne. 2014. <em>An Indigenous Peoples\u2019 History of the United States<\/em>. Boston: Beacon.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 15.1101684570312pt;text-indent: 0pt\">El-Zaher, Sumaya. 2018. \u201cThe Father of the Theory of Evolution: Al-Jahiz and His Book of Animals.\u201d MVSLIM.com, October 9. Accessed August 27, 2022. https:\/\/mvslim.com\/the-father-of-the-theory-of-evolution-al-jahiz-and-his-book-of-animals\/.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Elshakry, Marwa. 2010. \u201cWhen Science Became Western: Historiographical Reflections.\u201d <em>ISIS<\/em> 101 (1). Accessed November 20, 2022. <a class=\"rId107\" href=\"https:\/\/www.journals.uchicago.edu\/doi\/10.1086\/652691\">https:\/\/www.journals.uchicago.edu\/doi\/10.1086\/652691<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Gladwell, Malcolm. 2007. <em>Blink: The Power of Thinking without Thinking<\/em>. New York: Back Bay<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Guadagno, Rosanna E., Bradley M. Okdie, and Sara A. Kruse. 2012. \u201cDating Deception: Gender, Online Dating, and Exaggerated Self-Presentation.\u201d <em>Computers in Human Behavior<\/em> 28 (2): 642\u2013647.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 47.9759521484375pt;text-indent: 0pt\">Harmon, Katherine. 2011. \u201cEvolution Abroad: Creationism Evolves in Science Classrooms around the Globe.\u201d <em>Scientific American, <\/em>March 3. Accessed May 25, 2022. https:\/\/www.scientificamerican.com\/article\/evolution-education-abroad\/.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 47.9759521484375pt;text-indent: 0pt\">Harrison, Ellie. 2018. \u201cThis is What the Perfect Body Looks Like - According to Science.\u201d <em>Radiotimes.com<\/em>. Accessed June 14, 2023. https:\/\/www.radiotimes.com\/tv\/documentaries\/this-is-what-the-perfect-body-looks-like-according-to-science\/.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Hatton, Tim. 2014. \u201cWhy Did Humans Grow Four Inches in 100 Years? It Wasn\u2019t Just Diet.\u201d <em>The Conversation<\/em>, May 1. https:\/\/theconversation.com\/why-did-humans-grow-four-inches-In-100-years-it-wasnt-just-diet-25919.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Hays, J. N. 2005. <em>Epidemics and Pandemics: Their Impacts on Human History<\/em>. Santa Barbara, CA: ABC-CLIO.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Hutton, James. 1788. <em>Theory of the Earth<\/em>. Transactions of the Royal Society of Edinburgh, Volume 1. Scotland: Royal Society of Edinburgh.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 2.458251953125pt;text-indent: 0pt\">IPCC. 2022. <em>Climate Change 2022: Impacts, Adaptation, and Vulnerability. <\/em>Contribution of Working Group II to the Sixth Assessment Report of the Intergovernmental Panel on Climate Change, edited by H.-O. P\u00f6rtner, D.C. Roberts, M. Tignor, E.S. Poloczanska, K. Mintenbeck, A. Alegr\u00eda, M. Craig, S. Langsdorf, S. L\u00f6schke, V. M\u00f6ller, A. Okem, and B. Rama. Cambridge: Cambridge University Press. https:\/\/www.ipcc.ch\/report\/ar6\/wg2\/.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 2.458251953125pt;text-indent: 0pt\">Jones, Stephen. 2017. \u201cReligion, Science, and Evolutionary Theory\u201d[an interview with Salman Hameed]. Podcast, <em>The Religious Studies Project<\/em>, January 30. Accessed May 29, 2023. https:\/\/www.religiousstudiesproject.com\/podcast\/religion-science-and-evolutionary-theory\/.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 2.458251953125pt;text-indent: 0pt\">Kutschera, Ulrich, and Karl J. Niklas. 2004. \u201cThe Modern Theory of Biological Evolution: An Expanded Synthesis.\u201d <em>Naturwissenschaften<\/em> 91: 255\u2013276.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 2.458251953125pt;text-indent: 0pt\">Leclerc, Georges-Louis, Comte de Buffon. 1749-1804. <em>Histoire Naturelle<\/em>. Volumes 1-36. Paris: <em>Imprimerie Royale <\/em>(Royal Printing House).<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 31.8324584960938pt;text-indent: 0pt\">Lents, Nathan H. 2018. <em>Human Errors: A Panorama of Our Glitches, from Pointless Bones to Broken Genes. <\/em>Boston: Houghton Mifflin Harcourt.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 31.8324584960938pt;text-indent: 0pt\">Lerner, Lawrence S. 2000. <em>Good Science, Bad Science: Teaching Evolution in the States<\/em>. Washington, DC: Thomas B. Fordham Foundation.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Lieberman, Daniel E. 2011. <em>The Evolution of the Human Head<\/em>. Cambridge, MA: Harvard University Press.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 15.0872802734375pt;text-indent: 0pt\">Lindberg, David C. 1992. <em>The Beginnings of Western Science: The European Scientific Tradition in Philosophical, Religious, and Institutional Context, 600 B.C. to A.D. 1450<\/em>. Chicago: The University of Chicago Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Linnaeus, Carl. 1736. <em>Systema Naturae<\/em>. First Edition. Stockholm: Laurentius Salvius.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 21.6511840820312pt;text-indent: 0pt\">Love, Shayla. 2020. \u201cA Thousand Years before Darwin, Islamic Scholars Were Writing about Evolution.\u201d VICE World News, October 5. Accessed August 27, 2022. https:\/\/www.vice.com\/en\/article\/ep4ykn\/a-thousand-years-before-darwin-islamic-scholars-were-writing-about-natural-selection.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Main, Douglas. 2013. \u201cAncient Mutation Explains Missing Wisdom Teeth.\u201d <em>Live Science<\/em>, March 13. https:\/\/www.livescience.com\/27529-missing-wisdom-teeth.html.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Malthus, Thomas Robert. 1798. <em>An Essay on the Principle of Population<\/em>. London: J. Johnson.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Masci, David. 2019. \u201cFor Darwin Day, 6 Facts About the Evolution Debate.\u201d Pew Research Center. Accessed June 14, 2023. https:\/\/www.pewresearch.org\/short-reads\/2019\/02\/11\/darwin-day\/.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Masood, Ehsan. 2009. \u201cIslam\u2019s Evolutionary Legacy.\u201d <em>The Guardian<\/em>, March 1. Accessed February 13, 2023. https:\/\/www.theguardian.com\/commentisfree\/belief\/2009\/feb\/27\/islam-religion-evolution-science.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Miller, Jon D., Eugenie C. Scott, Mark S. Ackerman, Belen Laspra, Glenn Branch, Carmelo Polino, and Jordan S. Huffaker. 2022. \u201cPublic Acceptance of Evolution in the United States, 1985\u20132020.\u201d <em>Public Understanding of Science<\/em> 31 (2): 223\u2013238.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Miller, Jon D., Eugenie C. Scott, and Shinji Okamoto. 2006, August 11. \u201cPublic Acceptance of Evolution.\u201d <em>Science<\/em> 313 (5788): 765-766. <a class=\"rId108\" href=\"https:\/\/doi.org\/10.1126\/science.1126746\">DOI: 10.1126\/science.1126746<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 46.4606018066406pt;text-indent: 0pt\">Moore, John A. 1993. <em>Science as a Way of Knowing: The Foundations of Modern Biology<\/em>. Cambridge, MA: Harvard University Press.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 46.4606018066406pt;text-indent: 0pt\">Moore, Michael P., Kaitlyn Hersch, Chanont Sricharoen, Sarah Lee, Caitlin Reice, Paul Rice, Sophie Kronick, Kim A. Medley, and Kasey D. Fowler-Finn. 2021. \u201cSex-Specific Ornament Evolution Is a Consistent Feature of Climatic Adaptation across Space and Time in Dragonflies.\u201d <em>PNAS <\/em>118 (28): e2101458118. https:\/\/doi.org\/10.1073\/pnas.210145818.<\/p>\n<p class=\"import-Normal\">National Public Radio. 2023. \u201cForest Lizards Have Genetically Morphed to Survive Life in the City, Researchers Say.\u201d <em>National Public Radio (NPR), <\/em>January 10. Accessed February 19, 2023. https:\/\/www.npr.org\/2023\/01\/10\/1148150056\/forest-lizards-genetically-morphed-cities.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 46.4606018066406pt;text-indent: 0pt\">National Science Foundation. 2023. \u201cUrban Lizards Share Genomic Markers Not Found in Forest-Dwellers.\u201d National Science Foundation, February 7. Accessed May 25, 2023. <a class=\"rId109\" href=\"https:\/\/beta.nsf.gov\/news\/urban-lizards-share-genomic-markers-not-found\">https:\/\/beta.nsf.gov\/news\/urban-lizards-share-genomic-markers-not-found<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Novembre, John, Alison P. Galvani, and Montgomery Slatkin. 2005. \u201cThe Geographic Spread of the CCR5 \u039432 HIV-Resistance Allele.\u201d <em>PLoS Biology<\/em> 3 (11): e339.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Pamuk, \u015eevket. 2007. \u201cThe Black Death and the Origins of the \u2018Great Divergence\u2019 across Europe, 1300\u20131600.\u201d <em>European Review of Economic History<\/em> 11 (3): 289\u2013317.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 26.6983642578125pt;text-indent: 0pt\">Paterlini, Marta. 2007. \u201cThere Shall Be Order: The Legacy of Linnaeus in the Age of Molecular Biology.\u201d <em>EMBO Reports<\/em> 8 (9): 814\u2013816. <a class=\"rId110\" href=\"https:\/\/doi.org\/10.1038\/sj.embor.7401061\">https:\/\/doi.org\/10.1038\/sj.embor.7401061<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 26.6983642578125pt;text-indent: 0pt\">Pew Research Center. 2019. \u201cDarwin in America: The Evolution Debate in the United States.\u201d Pew Research Center, February 6, 2019. Accessed June 13, 2023. https:\/\/www.pewresearch.org\/religion\/2019\/02\/06\/darwin-in-america-2\/.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 35.9031982421875pt;text-indent: 0pt\">Pew Research Center. 2014. \u201cFighting Over Darwin, State by State.\u201d Pew Research Center, February 4, 2009; updated February 3, 2014. Accessed May 25, 2022. <a class=\"rId111\" href=\"https:\/\/www.pewresearch.org\/religion\/2009\/02\/04\/fighting-over-darwin-state-by-state\/\">https:\/\/www.pewresearch.org\/religion\/2009\/02\/04\/fighting-over-darwin-state-by-state\/<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 35.9031982421875pt;text-indent: 0pt\">Plutynski, Anya. 2009. \u201cThe Modern Synthesis.\u201d <em>Routledge Encyclopedia of Philosophy<\/em>, November 15 (updated December 14, 2009). Accessed November 27, 2022. https:\/\/philsci-archive.pitt.edu\/15335\/.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 15.7255249023438pt;text-indent: 0pt\">Plutzer, Eric, Glenn Branch, and Ann Reid. 2020. \u201cTeaching Evolution in the U.S. Public Schools: A Continuing Challenge.\u201d <em>Evolution: Education and Outreach<\/em>13: Article 14. https:\/\/evolution-outreach.biomedcentral.com\/articles\/10.1186\/s12052-020-00126-8.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 15.7255249023438pt;text-indent: 0pt\">Proceedings of the National Academy of Sciences (PNAS). 2023. \u201cHow Lizards Adapt to Urban Living.\u201d <em>Science Sessions Podcast, <\/em>February 13, . Accessed February 19, 2023. https:\/\/www.pnas.org\/post\/podcast\/lizards-adapt-urban-living.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 15.7255249023438pt;text-indent: 0pt\">Public Broadcasting System. 2001. \u201cGeorges Cuvier.\u201d <em>WGBH Evolution Library<\/em>. Accessed May 26, 2022. https:\/\/www.pbs.org\/wgbh\/evolution\/library\/02\/1\/l_021_01.html# :~:text=With%20elegant%20studies%20of%20the,as%20incontrovertible%20proof%20of %20extinctions.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 15.7255249023438pt;text-indent: 0pt\">Ray, John. 1686-1704. <em>Historia plantarum<\/em>. London: Clark. Volume 1 (1686), Volume II (1688), Volume III (1704).<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 15.7255249023438pt;text-indent: 0pt\">Richards, Richard A. 1998. \u201cDarwin, Domestic Breeding, and Artificial Selection.\u201d <em>Endeavour<\/em> 22 (3): 106\u2013109.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0.891006469726562pt;margin-right: 12.08837890625pt;text-indent: 0.648994445800781pt\">Solloch, Ute V., Kathrin Lang, Vinzenz Lange, and Irena B\u00f6hme. 2017. \u201cFrequencies of Gene Variant CCR5-\u039432 in 87 Countries Based on Next-Generation Sequencing of 1.3 Million Individuals Sampled from 3 National DKMS Donor Centers.\u201d <em>Human Immunology<\/em> 78 (11\u201312): 701-717.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 26.1246337890625pt;text-indent: 0pt\">Stewart, Thomas A., Justin B. Lemberg, Ailis Daly, Edward B. Daeschler, and Neil H. Shubin. 2022. \u201cA New Epistostegalian from the Late Devonian of the Canadian Arctic.\u201d <em>Nature<\/em> 608 (7923): 563\u2013568.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 14.37353515625pt;text-indent: 0pt\">Tasci, Ufuk Necat. 2020. \u201cHow a 10-Century Muslim Physicist Discovered How Humans See.\u201d <em>T<\/em><em>RT World<\/em>, May 25. Accessed May 18, 2022. https:\/\/www.trtworld.com\/magazine\/how-a-10th-century-muslim-physicist-discovered-how-humans-see-36620.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Tbakhi, Abdelghani, and Samir S. Amr. 2007. \u201cIbn Al-Haytham: Father of Modern Optics.\u201d <em>Annals of Saudi Medicine<\/em> 27 (6): 464\u2013467. UNAIDS. 2021. \u201cGlobal HIV &amp; AIDS Statistics\u2014Fact Sheet.\u201d https:\/\/www.unaids.org\/en\/resources\/fact-sheet.<\/p>\n<p class=\"import-Normal\" style=\"margin-right: 48.1709594726562pt\">UNESCO.org. 2015. \u201cInternational Year of Light: Ibn al Haytham, Pioneer of Modern Optics celebrated at UNESCO<em>.<\/em>\u201d UNESCO, September 8. Accessed May 18, 2022. https:\/\/www.unesco.org\/en\/articles\/international-year-light-ibn-al-haytham-pioneer-modern-optics-celebrated-unesco.<\/p>\n<p class=\"import-Normal\" style=\"margin-right: 48.1709594726562pt\">University of California Berkeley Museum of Paleontology. N.d. \u201cThe History of Evolutionary Thought\u2014Uniformitarianism: Charles Lyell.\u201d <em>Understanding Evolution <\/em>website. Accessed February 13, 2023. https:\/\/evolution.berkeley.edu\/the-history-of-evolutionary-thought\/1800s\/uniformitarianism-charles-lyell\/.<\/p>\n<p class=\"import-Normal\" style=\"margin-right: 48.1709594726562pt\">University of Cambridge. N.d. \u201cDarwin and Dogs.\u201d <em>The Darwin Correspondence Project <\/em>website. Accessed February 17, 2023. https:\/\/www.darwinproject.ac.uk\/commentary\/curious\/darwin-and-dogs.<\/p>\n<p class=\"import-Normal\" style=\"margin-right: 48.1709594726562pt\">Urbach, Peter Michael, Anthony M. Quinton, and Kathleen Marguerite Lea. \u201cFrancis Bacon: British Author, Philosopher, and Statesman.\u201d <em>Encyclopaedia Britannica<\/em>. Last updated May 12, 2023. https:\/\/www.britannica.com\/biography\/Francis-Bacon-Viscount-Saint-Alban.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 17.7549438476562pt;text-indent: 0pt\">Ward, Peter. 2018. <em>Lamarck\u2019s Revenge: How Epigenetics Is Revolutionizing Our Understanding<\/em> <em>of Evolution\u2019s Past and Present. <\/em>New York: Bloomsbury.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 17.7549438476562pt;text-indent: 0pt\">Weismann, August. 1892. <em>Das Keimplasma: Eine Theorie der Vererbung<\/em> (<em>The Germ Plasm: a Theory of Inheritance<\/em>). Jena (Germany): Fischer.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 17.7549438476562pt;text-indent: 0pt\">Weismann, August. 1889. Translations. <em>Essays upon Heredity<\/em>. Oxford: Clarendon. Accessed November 27, 2022. E-copy available at https:\/\/www.esp.org\/books\/weismann\/essays\/facsimile\/.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 17.7549438476562pt;text-indent: 0pt\">Winchell, Kristen M., Shane C. Campbell-Staton, Jonathan B. Losos, and Anthony Geneva. 2023. \u201cGenome-Wide Parallelism Underlies Contemporary Adaptation In Urban Lizards.\u201d <em>PNAS<\/em> 120 (3): e2216789120. https:\/\/doi.org\/10.1073\/pnas.2216789120.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 17.7549438476562pt;text-indent: 0pt\">Wood, Bernard. 2005. <em>Human Evolution: A Very Short Introduction<\/em>. Oxford: Oxford University Press.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;margin-left: 0pt;margin-right: 17.7549438476562pt;text-indent: 0pt\">Zou, Yawen. 2015. \"The Germ-Plasm: a Theory of Heredity (1893), by August Weismann.\" <em>Embryo Project Encyclopedia<\/em>, January 26. 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