{"id":1683,"date":"2026-08-20T17:15:39","date_gmt":"2026-08-20T21:15:39","guid":{"rendered":"https:\/\/opentextbooks.concordia.ca\/explorations3\/?post_type=chapter&#038;p=1683"},"modified":"2026-09-01T13:36:07","modified_gmt":"2026-09-01T17:36:07","slug":"primate-evolution-test-wip","status":"publish","type":"chapter","link":"https:\/\/opentextbooks.concordia.ca\/explorations3\/chapter\/primate-evolution-test-wip\/","title":{"raw":"Primate Evolution","rendered":"Primate Evolution"},"content":{"raw":"<div class=\"__UNKNOWN__\">\r\n<div class=\"textbox\">\r\n\r\nLearn more about <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/about-the-authors\/\">authors and editors<\/a>,<a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/student-contributors\/\"> Hess' student contributions<\/a>, and <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/versioning-history\/\">versioning history<\/a>.\r\n<div class=\"textbox textbox--learning-objectives\"><header class=\"textbox__header\">\r\n<h2 class=\"textbox__title\">Learning Objectives<\/h2>\r\n<\/header>\r\n<div class=\"textbox__content\">\r\n<ul>\r\n \t<li>Understand the major trends in primate evolution from the origin of primates to the origin of our own species.<\/li>\r\n \t<li>Learn about primate adaptations and how they characterize major primate groups.<\/li>\r\n \t<li>Discuss the kinds of evidence that anthropologists use to find out how extinct primates are related to each other and to living primates.<\/li>\r\n \t<li>Recognize how the changing geography and climate of Earth have influenced where and when primates have thrived or gone extinct.<\/li>\r\n<\/ul>\r\n<\/div>\r\n<\/div>\r\n<p class=\"import-Normal\">The first fifty million years of primate evolution was a series of <strong>[pb_glossary id=\"1683\"]adaptive radiations[\/pb_glossary]<\/strong> leading to the diversification of the earliest lemurs, monkeys, and apes. The primate story begins in the canopy and understory of conifer-dominated forests, with our small, furtive ancestors subsisting at night, beneath the notice of day-active dinosaurs.<\/p>\r\n<p class=\"import-Normal\">From the ancient [pb_glossary id=\"1684\"]<strong>plesiadapiforms<\/strong>[\/pb_glossary] (archaic primates) to the earliest groups of true primates ([pb_glossary id=\"1686\"]<strong>euprimates<\/strong>[\/pb_glossary]) (Bloch and Boyer 2002), the origin of our own order is characterized by the struggle for new food sources and microhabitats in the arboreal setting. Climate change forced major extinctions as the northern continents became increasingly dry, cold, and seasonal and as tropical rainforests gave way to deciduous forests, woodlands, and eventually grasslands. Lemurs, lorises, and tarsiers\u2014once diverse groups containing many species\u2014became rare, except for lemurs in Madagascar, where there were no anthropoid competitors and perhaps few predators. Meanwhile, <strong>[pb_glossary id=\"1685\"]anthropoids[\/pb_glossary]<\/strong> (monkeys and apes) likely emerged in Asia and then dispersed across parts of the northern hemisphere, Africa, and ultimately South America. The movement of continents, shifting sea levels, and changing patterns of rainfall and vegetation contributed to the developing landscape of primate biogeography, morphology, and behaviour. Today\u2019s primates provide modest reminders of the past diversity and remarkable adaptations of their extinct relatives. This chapter explores the major trends in primate evolution from the origin of the Order Primates to the beginnings of our own lineage, providing a window into these stories from our ancient past.<\/p>\r\n\r\n<h2 class=\"import-Normal\">Major Hypotheses About Primate Origins<\/h2>\r\n<p class=\"import-Normal\">For many groups of mammals, there is a key feature that led to their success. A good example is powered flight in bats. Primates lack a feature like this (see Chapter 6). Instead, if there is something unique about primates, it is probably a group of features rather than one single thing. Because of this, anthropologists and palaeontologists struggle to describe an ecological scenario that could explain the rise and success of our own order. Three major hypotheses have been advanced to consider the origin of primates and to explain what makes our order distinct among mammals (Figure 9.1); these are described below.<\/p>\r\n\r\n\r\n[caption id=\"attachment_277\" align=\"aligncenter\" width=\"634\"]<img class=\"wp-image-255\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2023\/03\/8.1.jpg\" alt=\"Primates swinging in tree, eating an insect, and eating fruit.\" width=\"634\" height=\"221\" \/> Figure 9.1: The three major hypotheses are (a) the arboreal hypothesis, (b) the visual predation hypothesis, and (c) the angiosperm-primate coevolution hypothesis. Credit: Primate origin hypotheses original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by <a class=\"rId13\" href=\"https:\/\/marynelsonstudio.com\">Mary Nelson<\/a> is under a <a class=\"rId14\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.[\/caption]\r\n<h3 class=\"import-Normal\"><strong>Arboreal Hypothesis<\/strong><\/h3>\r\n<p class=\"import-Normal\">In the 1800s, many anthropologists viewed all animals in relation to humans. That is, animals that were more like humans were considered to be more \u201cadvanced\u201d and those lacking humanlike features were considered more \u201cprimitive.\u201d This way of thinking was particularly obvious in studies of primates. A more modern way of referring to members of a group that lack certain evolutionary innovations seen in other members is to call them [pb_glossary id=\"1688\"]<strong>plesiomorphic<\/strong>[\/pb_glossary] (literally \u201canciently shaped\u201d). The state of their morphological features is sometimes referred to as [pb_glossary id=\"1689\"]<strong>ancestral<\/strong><strong> traits<\/strong>[\/pb_glossary].<\/p>\r\n<p class=\"import-Normal\">Thus, when anthropologists sought features that separate primates from other mammals, they focused on features that were least developed in lemurs and lorises, more developed in monkeys, and most developed in apes (Figure 9.2). Frederic Wood Jones, one of the leading anatomist-anthropologists of the early 1900s, is usually credited with the Arboreal Hypothesis of primate origins (Jones 1916). This hypothesis holds that many of the features of primates evolved to improve locomotion in the trees; this way of getting around is referred to as arboreal. For example, the grasping hands and feet of primates are well suited to gripping tree branches of various sizes and our flexible joints are good for reorienting the extremities in many different ways. A mentor of Jones, Grafton Elliot Smith, had suggested that the reduced olfactory system, acute vision, and forward-facing eyes of primates are adaptations for making accurate leaps and bounds through a complex, three-dimensional canopy (Smith 1912). The forward orientation of the eyes in primates causes the visual fields to overlap, enhancing depth perception, especially at close range. Evidence to support this hypothesis includes the facts that many extant primates are arboreal, and the plesiomorphic members of most primate groups are dedicated arborealists. The Arboreal Hypothesis was well accepted by most anthropologists at the time and for decades afterward.<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"663\"]<img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image27-2.png\" alt=\"Diagram shows primates descended from Plesiadapiforms.\" width=\"663\" height=\"543\" \/> Figure 9.2: Primate family tree showing major groups. Disconnected lines show uncertainty about relationships. Two lines lead to tarsiers from different possible groups of origin. <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 class=\"rId16\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Primate family tree (Figure 8.2)<\/a> by Jonathan M. G. Perry 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<h3 class=\"import-Normal\"><strong>Visual Predation Hypothesis<\/strong><\/h3>\r\n<p class=\"import-Normal\">In the late 1960s and early 1970s, Matt Cartmill studied and tested the idea that the characteristic features of primates evolved in the context of arboreal locomotion. Cartmill noted that squirrels climb trees (and even vertical walls) very effectively, even though they lack some of the key adaptations of primates. As members of the Order Rodentia, squirrels also lack the hand and foot anatomy of primates. They have claws instead of flattened nails and their eyes face more laterally than those of primates. Cartmill reasoned that there must be some other explanation for the unique traits of primates. He noted that some nonarboreal animals share at least some of these traits with primates; for example, cats and predatory birds have forward-facing eyes that enable visual field overlap. Cartmill suggested that the unique suite of features in primates is an adaptation to detecting insect prey and guiding the hands (or feet) to catch insects (Cartmill 1972). His hypothesis emphasizes the primary role of vision in prey detection and capture; it is explicitly comparative, relying on form-function relationships in other mammals and nonmammalian vertebrates. According to Cartmill, many of the key features of primates evolved for preying on insects in this special manner (Cartmill 1974).<\/p>\r\n\r\n<h3 class=\"import-Normal\"><strong>Angiosperm-Primate Coevolution Hypothesis<\/strong><\/h3>\r\n<p class=\"import-Normal\">The visual predation hypothesis was unpopular with some anthropologists. One reason for this is that many primates today are not especially predatory. Another is that, whereas primates do seem well adapted to moving around in the smallest, terminal branches of trees, insects are not necessarily easier to find there. A counterargument to the visual predation hypothesis is the angiosperm-primate coevolution hypothesis. Primate ecologist Robert Sussman (Sussman 1991) argued that the few primates that eat mostly insects often catch their prey on the ground rather than in tree branches. Furthermore, predatory primates often use their ears more than their eyes to detect prey. Finally, most early primate fossils show signs of having been omnivorous rather than insectivorous. Instead, he argued, the earliest primates were probably seeking fruit. Fruit (and flowers) of angiosperms (flowering plants) often develop in the terminal branches. Therefore, any mammal trying to access those fruits must possess anatomical traits that allow them to maintain their hold on thin branches and avoid falling while reaching for the fruits. Primates likely evolved their distinctive visual traits and extremities in the Paleocene (approximately 65 million to 54 million years ago) and Eocene (approximately 54 million to 34 million years ago) epochs, just when angiosperms were going through a revolution of their own\u2014the evolution of large, fleshy fruit that would have been attractive to a small arboreal mammal. Sussman argued that, just as primates were evolving anatomical traits that made them more efficient fruit foragers, angiosperms were also evolving fruit that would be more attractive to primates to promote better seed dispersal. This mutually beneficial relationship between the angiosperms and the primates was termed coevolution or more specifically [pb_glossary id=\"1691\"]<strong>diffuse coevolution<\/strong>.[\/pb_glossary]<\/p>\r\n<p class=\"import-Normal\">At about the same time, D. Tab Rasmussen noted several parallel traits in primates and the South American woolly opossum, <em>Caluromys<\/em>. He argued that early primates were probably foraging on both fruits and insects (Rasmussen 1990). As is true of <em>Caluromys<\/em> today, early primates probably foraged for fruits in the terminal branches of angiosperms, and they probably used their visual sense to aid in catching insects. Insects are also attracted to fruit (and flowers), so these insects represent a convenient opportunity for a primarily fruit-eating primate to gather protein. This solution is a compromise between the visual predation hypothesis and the angiosperm-primate coevolution hypothesis. It is worth noting that other models of primate origins have been proposed, and these include the possibility that no single ecological scenario can account for the origin of primates.<\/p>\r\n\r\n<h2 class=\"import-Normal\">The Origins of Primates<\/h2>\r\n<h3 class=\"import-Normal\"><strong>Paleocene: Mammals in the Wake of Dinosaur Extinctions<\/strong><\/h3>\r\n<p class=\"import-Normal\">Placental mammals, including primates, originated in the Mesozoic Era (approximately 251 million to 65.5 million years ago), the Age of Dinosaurs. During this time, most placental mammals were small, probably nocturnal, and probably avoided predators via camouflage and slow, quiet movement. It has been suggested that the success and diversity of the dinosaurs constituted a kind of ecological barrier to Mesozoic mammals. The extinction of the dinosaurs (and many other organisms) at the end of the Cretaceous Period (approximately 145.5\u201365.5 million years ago) might have opened up these ecological niches, leading to the increased diversity and disparity in mammals of the Tertiary Period (approximately 65.5\u20132.5 million years ago).<\/p>\r\n<p class=\"import-Normal\">The Paleocene was the first epoch in the Age of Mammals. Soon after the Cretaceous-Tertiary (K-T) extinction event, new groups of placental mammals appear in the fossil record. Many of these groups achieved a broad range of sizes and lifestyles as well as a great number of species before declining sometime in the Eocene (or soon thereafter). These groups were ultimately replaced by the modern orders of placental mammals (Figure 9.3). It is unknown whether these replacements occurred gradually, for example by competitive exclusion, or rapidly, perhaps by sudden geographic dispersals with replacement. In some senses, the Paleocene might have been a time of recovery from the extinction event; it was cooler and more seasonal globally than the subsequent Eocene.<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"628\"]<img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image26.jpg\" alt=\"Person in front of a mural depicting forest animals.\" width=\"628\" height=\"511\" \/> Figure 9.3: A mural of Eocene flora and fauna in North America. Credit: <a class=\"rId19\" href=\"https:\/\/flickr.com\/photos\/126377022@N07\/18404106406\">Image from page 27 of \"Annual report for the year ended June 30 ...\" (1951)<\/a> by <a class=\"rId20\" href=\"https:\/\/flickr.com\/photos\/internetarchivebookimages\/\">Internet Archive Book Images<\/a> has been designated to the <a class=\"rId21\" href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/\">public domain (CC0)<\/a>. This photograph of the mural \"Fauna and flora of middle Eocene in the Wyoming area\" by Jay Matternes, was originally published by the <a class=\"rId22\" href=\"https:\/\/www.si.edu\/\">Smithsonian<\/a>, and can be viewed in context in the <a class=\"rId23\" href=\"https:\/\/archive.org\/details\/annualreportfory1961united\/page\/7\/mode\/1up?view=theater\">online version of this book<\/a>.[\/caption]\r\n<h3 class=\"import-Normal\"><strong>Plesiadapiforms, the Archaic Primates<\/strong><\/h3>\r\n<p class=\"import-Normal\">The Paleocene epoch saw the emergence of several families of mammals that have been implicated in the origin of primates. These are the plesiadapiforms, which are archaic primates, meaning they possessed some primate features and lacked others. The word <em>plesiadapiform <\/em>means \u201calmost adapiform,\u201d a reference to some similarities between some plesiadapiforms and some adapiforms (or adapoids; later-appearing true primates)\u2014mainly in the molar teeth. Because enamel fossilizes better than other parts of the body, the molar teeth are the parts most often found and first discovered for any new species. Thus, dental similarities were often the first to be noticed by early mammalian paleontologists, partly explaining why plesiadapiforms were thought to be primates. Major morphological differences between plesidapiforms and euprimates (true primates) were observed later when more parts of plesiadapiform skeletons were discovered. Many plesiadapiforms have unusual anterior teeth and most have digits possessing claws rather than nails. So far, no plesiadapiform ever discovered has a postorbital bar (seen in extant <strong>[pb_glossary id=\"1712\"]strepsirrhines[\/pb_glossary]<\/strong>) or septum (as seen in <strong>[pb_glossary id=\"1713\"]haplorhines[\/pb_glossary]<\/strong>), and whether or not the <strong>[pb_glossary id=\"1711\"]auditory bulla[\/pb_glossary]<\/strong> was formed by the [pb_glossary id=\"1714\"]<strong>petrosal bone<\/strong> [\/pb_glossary]remains unclear for many plesiadapiform specimens. Nevertheless, there are compelling reasons (partly from new skeletal material) for including plesiadapiforms within the Order Primates.<\/p>\r\n\r\n<h4 class=\"import-Normal\"><em>Geographic and Temporal Distribution<\/em><\/h4>\r\n<p class=\"import-Normal\"><em>Purgatorius<\/em> is generally considered to be the earliest primate. This Paleocene mammal is known from teeth that are very plesiomorphic for a primate. It has some characteristics that suggest it is a basal plesiadapiform, but there is very little to link it specifically with euprimates (see Clemens 2004). Its ankle bones suggest a high degree of mobility, signaling an arboreal lifestyle (Chester Et al. 2015). <em>Purgatorius<\/em> is plesiomorphic enough to have given rise to all primates, including the plesiadapiforms. However, new finds suggest that this genus was more diverse and had more differing tooth morphologies than previously appreciated (Wilson Mantilla Et al. 2021). Plesiadapiform families were numerous and diverse during parts of the Paleocene in western North America and western Europe, with some genera (e.g., <em>Plesiadapis<\/em>; see Figure 9.4) living on both continents (Figure 9.5). Thus, there were probably corridors for plesiadapiform dispersal between the two continents, and it stands to reason that these mammals were living all across North America, including in the eastern half of the continent and at high latitudes. A few plesiadapiforms have been described from Asia (e.g., <em>Carpocristes<\/em>), but the affinities of these remain uncertain.<\/p>\r\n\r\n<div style=\"text-align: left\">\r\n<table class=\"aligncenter\" style=\"width: 473.25pt\"><caption>Figure 9.4: Families of plesiadapiforms with example genera and traits: a table. Credit: Plesiadapiforms table original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Jonathan M. G. Perry and Stephanie L. Canington is under a <a class=\"rId24\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Content derived from Fleagle 2013.<\/caption>\r\n<thead>\r\n<tr style=\"height: 25pt\">\r\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Family<\/strong><\/p>\r\n&nbsp;<\/td>\r\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Genera<\/strong><\/p>\r\n&nbsp;<\/td>\r\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Morphology<\/strong><\/p>\r\n&nbsp;<\/td>\r\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Location<\/strong><\/p>\r\n&nbsp;<\/td>\r\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Age<\/strong><sup><strong>1<\/strong><\/sup><\/p>\r\n&nbsp;<\/td>\r\n<\/tr>\r\n<\/thead>\r\n<tbody>\r\n<tr class=\"Table1-R\" style=\"height: 17pt\">\r\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Paromomyidae<\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\"><em>Ignacius<\/em><\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Long, dagger-like, lower incisor.<\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">North America and Europe<\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Early Paleocene to Late Eocene<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table1-R\" style=\"height: 18pt\">\r\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Carpolestidae<\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\"><em>Carpolestes<\/em><\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Plagiaulacoid dentition. Limb adaptations to terminal branch feeding. Grasping big toe.<\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">North America, Europe, and Asia<\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Middle Paleocene to Early Eocene<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table1-R\" style=\"height: 16pt\">\r\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Plesiadapidae<\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\"><em>Plesiadapis<\/em><\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Mitten-like upper incisor. Diastema. Large body size for group.<\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">North America and Europe<\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Middle Paleocene to Early Eocene<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table1-R\" style=\"height: 1pt\">\r\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\" colspan=\"4\">\r\n<p class=\"import-Normal\"><sup>1<\/sup> Derived from Fleagle 2013.<\/p>\r\n<\/td>\r\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\">\r\n<p class=\"import-Normal\"><\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr>\r\n<td><\/td>\r\n<td><\/td>\r\n<td><\/td>\r\n<td><\/td>\r\n<td><\/td>\r\n<\/tr>\r\n<\/tbody>\r\n<\/table>\r\n<\/div>\r\n\r\n[caption id=\"attachment_277\" align=\"aligncenter\" width=\"555\"]<img class=\"wp-image-258 \" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5-5-e1691791897574.png\" alt=\"Global map with not fully formed continents.\" width=\"555\" height=\"308\" \/> Figure 9.5: Map of the world in the Paleocene, highlighting plesiadapiform localities on lands that would become North America, southern Europe, and eastern Asia. Credit: <a class=\"rId26\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Paleocene Map with Plesiadapiform Localities (Figure 8.4)<\/a> original to<a class=\"rId27\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"> Expl<\/a><a class=\"rId28\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">orations: An Open Invitation to Biological Anthropology<\/a> by Elyssa Ebding at <a class=\"rId29\" href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\">GeoPlace, California State University, Chico<\/a> is under a <a class=\"rId30\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Localities based on Fleagle 2013, 211.[\/caption]\r\n<h4 class=\"import-Normal\"><em>General Morphological Features<\/em><\/h4>\r\n<p class=\"import-Normal\">Although there is much morphological variation among the families of plesiadapiforms, some common features unite the group. Most plesiadapiforms were small, the largest being about three kilograms (approximately 7 lbs.; <em>Plesiadapis cookei<\/em>). They had small brains and fairly large snouts, with eyes that faced more laterally than in euprimates. Many species show reduction and\/or loss of the canine and anterior premolars, with the resulting formation of a rodent-like <strong>[pb_glossary id=\"1172\"]diastema[\/pb_glossary] <\/strong>(a pronounced gap between the premolars and the incisors, with loss of at least the canine); this probably implies a herbivorous diet. Some families appear to have had very specialized diets, as suggested by unusual tooth and jaw shapes.<\/p>\r\n<p class=\"import-Normal\">Arguably the most interesting and unusual family of plesiadapiforms is the Carpolestidae. They are almost exclusively from North America (with a couple of possible members from Asia), and mainly from the Middle and Late Paleocene. Their molars are not very remarkable, being quite similar to those of some other plesiadapiforms (e.g., Plesiadapidae). However, their lower posterior premolars (p4) are laterally compressed and blade-like with vertical serrations topped by tiny cuspules. This unusual dental morphology is termed [pb_glossary id=\"1693\"]<strong><em>plagiaulacoid<\/em><\/strong> [\/pb_glossary] (Simpson 1933). The upper premolar occlusal surfaces are broad and are covered with many small cuspules; the blade-like lower premolar might have cut across these cuspules, between them, or both.<\/p>\r\n\r\n<div class=\"mceTemp\"><\/div>\r\n\r\n[caption id=\"\" align=\"alignleft\" width=\"314\"]<img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image1-5.png\" alt=\"Small brown animal with long tail.\" width=\"314\" height=\"178\" \/> Figure 9.6: An artistic rendition of Carpolestes simpsoni moving along a small diameter support. Credit: <a class=\"rId32\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:CarpolestesCL.png\">CarpolestesCL<\/a> by <a class=\"rId33\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Sisyphos23\">Sisyphos23<\/a> is under a <a class=\"rId34\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.[\/caption]\r\n<p class=\"import-Normal\">Many plesiadapiforms have robust limb bones with hallmarks of arboreality. Instead of having nails, most taxa had sharp claws on most or all of the digits. The extremities show grasping abilities comparable to those of primates and some arboreal marsupials. Nearly complete skeletons have yielded a tremendous wealth of information on locomotor and foraging habits. Many plesiadapiforms appear to have been able to cling to vertical substrates (like a broad tree trunk) using their sharp claws, propelling themselves upward using powerful hindlimbs, bounding along horizontal supports, grasping smaller branches, and moving head-first down tree trunks. In carpolestids in particular, the skeleton appears to have been especially well adapted to moving slowly and carefully in small terminal branches (Figure 9.6).<\/p>\r\n\r\n<\/div>\r\n<div class=\"textbox shaded\">\r\n<h3 class=\"import-Normal\">Dig Deeper: Debate on the Relationship of Plesiadapiforms to True Primates<\/h3>\r\n<p class=\"import-Normal\">In the middle of the twentieth century, treeshrews (Order Scandentia) were often considered part of the Order Primates, based on anatomical similarities between some treeshrews and primates. For many people, plesiadapiforms represented intermediates between primates and treeshrews, so plesiadapiforms were included in Primates as well.<\/p>\r\n<p class=\"import-Normal\">Studies of reproduction and brain anatomy in treeshrews and lemurs suggested that treeshrews are not primates (e.g., Martin 1968). This was soon followed by the suggestion to also expel plesiadapiforms (Martin 1972) from the Order Primates. Like treeshrews, plesiadapiforms lack a postorbital bar, nails, and details of the ear region that characterize true primates. Many paleoanthropologists were reluctant to accept this move to banish plesiadapiforms (e.g., F. S. Szalay, P. D. Gingerich).<\/p>\r\n<p class=\"import-Normal\">Later, K. Christopher Beard (1990) found that in some ways, the digits of paromomyid plesiadapiforms are actually more similar to those of dermopterans (Order Dermoptera), the closest living relatives of primates, than they are to those of primates themselves (but see Krause 1991). At the same time, Richard Kay and colleagues (1990) found that cranial circulation patterns and auditory bulla morphology in the paromomyid, <em>Ignacius <\/em>(see Figure 9.4), are more like those of dermopterans than of primates.<\/p>\r\n<p class=\"import-Normal\">For many anthropologists, this one-two punch effectively removed plesiadapiforms from the Order Primates. In the last two decades, the tide of opinion has turned again, with many researchers reinstating plesiadapiforms as members of the Order Primates. New and more complete specimens demonstrate that the postcranial skeletons of plesiadapiforms, including the hands and feet, were primate-like, not dermorpteran-like (Bloch and Boyer 2002, 2007). New fine-grained CT scans of relatively complete plesiadapiform skulls revealed that they share some key traits with primates to the exclusion of other placental mammals (Bloch and Silcox 2006). Most significant was the suggestion that <em>Carpolestes simpsoni <\/em>possessed an auditory bulla formed by the [pb_glossary id=\"1696\"]<strong>petrosal <\/strong><strong>bone<\/strong>[\/pb_glossary], like in all living primates.<\/p>\r\n<p class=\"import-Normal\">The debate about the status of plesiadapiforms continues, owing to a persistent lack of key bones in some species and owing to genuine complexity of the anatomical traits involved. Maybe plesiadapiforms were the ancestral stock from which all primates arose, with some plesiadapiforms (e.g., carpolestids) nearer to the primate <strong>[pb_glossary id=\"1723\"]stem[\/pb_glossary]<\/strong> than others.<\/p>\r\n\r\n<\/div>\r\n<div class=\"__UNKNOWN__\">\r\n<h3 class=\"import-Normal\"><strong>Adapoids and Omomyoids, the First True Primates<\/strong><\/h3>\r\n<h4 class=\"import-Normal\"><em>Geographic and Temporal Distribution<\/em><\/h4>\r\n<p class=\"import-Normal\">The first universally accepted fossil primates are the adapoids (Superfamily [pb_glossary id=\"1695\"]<strong>Adapoidea<\/strong>[\/pb_glossary]) and the omomyoids (Superfamily <strong>[pb_glossary id=\"1694\"]Omomyoidea[\/pb_glossary])<\/strong>. These groups become quite distinct over evolutionary time, filling mutually exclusive niches for the most part. However, the earliest adapoids are very similar to the earliest omomyoids.<\/p>\r\n<p class=\"import-Normal\">The adapoids were mainly diurnal and herbivorous, with some achieving larger sizes than any plesiadapiforms (10 kg; 22 lbs.). By contrast, the omomyoids were mainly nocturnal, insectivorous and frugivorous, and small.<\/p>\r\n<p class=\"import-Normal\">Both groups appear suddenly at the start of the Eocene, where they are present in western North America, western Europe, and India (Figure 9.7). This wide dispersal of early primates was probably due to the presence of rainforest corridors extending far into northern latitudes.<\/p>\r\n\r\n\r\n[caption id=\"attachment_277\" align=\"aligncenter\" width=\"539\"]<img class=\"wp-image-260\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image22-3-e1691792023503.png\" alt=\"Global map with not fully formed continents and omomyoid localities.\" width=\"539\" height=\"317\" \/> Figure 9.7: Map of the world in the Eocene, highlighting adapoid and omomyoid localities on lands that would become North America, southern Europe, Africa, and Asia. Credit: <a class=\"rId36\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Eocene Map with Adapoid and Omomyoid Localities (Figure 8.6)<\/a> original to <a class=\"rId37\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Elyssa Ebding at <a class=\"rId38\" href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\">GeoPlace, California State University, Chico<\/a> is under a <a class=\"rId39\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Localities based on Fleagle 2013, 229.[\/caption]\r\n<p class=\"import-Normal\">In North America and Europe, both groups achieved considerable diversity in the Middle Eocene, then mostly died out at the end of that epoch (Figure 9.8). In some Eocene rock formations in the western United States, adapoids and omomyoids make up a major part of the mammalian fauna. The Eocene of India has yielded a modest diversity of euprimates, some of which are so plesiomorphic that it is difficult to know whether they are adapoids or omomyoids (or even early anthropoids).<\/p>\r\n\r\n<div style=\"text-align: left\">\r\n<table class=\"aligncenter\" style=\"width: 473.25pt\"><caption>Figure 9.8: Families of adapoids and omomyoids with example genera and traits: a table. Credit: Adapoids and omomyoids table original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Jonathan M. G. Perry and Stephanie L. Canington is under a <a class=\"rId40\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Content derived from Fleagle 2013.<\/caption>\r\n<thead>\r\n<tr style=\"height: 25pt\">\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Family<\/strong><\/p>\r\n&nbsp;<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Genera<\/strong><\/p>\r\n&nbsp;<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Morphology<\/strong><\/p>\r\n&nbsp;<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Location<\/strong><\/p>\r\n&nbsp;<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Age<\/strong><sup><strong>1<\/strong><\/sup><\/p>\r\n&nbsp;<\/td>\r\n<\/tr>\r\n<\/thead>\r\n<tbody>\r\n<tr class=\"Table2-R\" style=\"height: 18pt\">\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Cercamoniidae<\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\"><em>Donrussellia<\/em><\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Variable in tooth number and jaw shape.<\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Europe and Asia<\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Early to Late Eocene<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table2-R\" style=\"height: 16pt\">\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Asiadapidae<sup>2<\/sup><\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\"><em>Asiadapis<\/em><\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Plesiomorphic teeth and jaw resemble early Omomyids.<\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Asia<\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Early Eocene<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table2-R\" style=\"height: 16pt\">\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Caenopithecidae<sup>3<\/sup><\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\"><em>Darwinius<\/em><\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Robust jaws with crested molars. Fewer premolars.<\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Europe, Africa, North America, and Asia<\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Middle to Late Eocene<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table2-R\" style=\"height: 16pt\">\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Adapidae<\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\"><em>Adapis<\/em><\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Fused mandible. Long molar crests. Large size and large chewing muscles.<\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Europe<\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Late Eocene to Early Oligocene<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table2-R\" style=\"height: 16pt\">\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Sivaladapidae<\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\"><em>Sivaladapis<\/em><\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Some large with robust jaws.<\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Asia<\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Middle Eocene to Late Miocene<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table2-R\" style=\"height: 16pt\">\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Notharctidae<sup>4<\/sup><\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\"><em>Notharctus<\/em><\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Canine sexual dimorphism. Lemur-like skull. Clinging and leaping adaptations.<\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">North America and Europe<\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Early to Middle Eocene<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table2-R\" style=\"height: 16pt\">\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Omomyidae<sup>5<\/sup><\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\"><em>Teilhardina<\/em><\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Small, nocturnal, frugivorous or insectivorous. Tarsier-like skull in some.<\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">North America, Europe, and Asia<\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Early Eocene to Early Miocene<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table2-R\" style=\"height: 16pt\">\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Microchoeridae<sup>6<\/sup><\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\"><em>Necrolemur<\/em><\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Long bony ear tubes. Tarsier-like lower limb adaptations for leaping.<\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Europe and Asia<\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Early Eocene to Early Oligocene<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table2-R\" style=\"height: 1pt\">\r\n<td class=\"Table2-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\" colspan=\"4\">\r\n<p class=\"import-Normal\"><sup>1<\/sup> Derived from Fleagle 2013.<\/p>\r\n<p class=\"import-Normal\"><sup>2<\/sup> See Dunn Et al. 2016 and Rose Et al. 2018.<\/p>\r\n<p class=\"import-Normal\"><sup>3<\/sup> See Kirk and Williams 2011 and Seiffert Et al. 2009.<\/p>\r\n<p class=\"import-Normal\"><sup>4<\/sup> See Gregory 1920.<\/p>\r\n<p class=\"import-Normal\"><sup>5<\/sup> See Beard and MacPhee 1994 and Strait 2001.<\/p>\r\n<p class=\"import-Normal\"><sup>6<\/sup> See Schmid 1979.<\/p>\r\n<\/td>\r\n<td class=\"Table2-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\">\r\n<p class=\"import-Normal\"><\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr>\r\n<td><\/td>\r\n<td><\/td>\r\n<td><\/td>\r\n<td><\/td>\r\n<td><\/td>\r\n<\/tr>\r\n<\/tbody>\r\n<\/table>\r\n<\/div>\r\n<p class=\"import-Normal\">Adapoids and omomyoids barely survived the Eocene-Oligocene extinctions, when colder temperatures, increased seasonality, and the retreat of rainforests to lower latitudes led to changes in mammalian biogeography. In North America, one genus (originally considered an omomyoid but recently reclassified as Adapoidea) persisted until the Miocene: <em>Ekgmowechashala<\/em> (Rose and Rensberger 1983). This taxon has highly unusual teeth and might have been a late immigrant to North America from Asia. In Asia, one family of adapoids, the Sivaladapidae, retained considerable diversity as late as the Late Miocene.<\/p>\r\n\r\n<h4 class=\"import-Normal\"><em>Adapoid Diversity<\/em><\/h4>\r\n<p class=\"import-Normal\">Adapoids were very diverse, particularly in the Eocene of North America and Europe. They can be divided into six families, with a few species of uncertain familial relationship. As a group, adapoids have some features in common, although much of what they share is plesiomorphic. Important features include the hallmarks of euprimates: postorbital bar, flattened nails, grasping extremities, and a petrosal bulla (Figures 9.9 and 9.10). In addition, some adapoids retain the ancestral dental formula of 2.1.4.3; that is, in each quadrant of the mouth, there are two incisors, one canine, four premolars, and three molars. In general, the incisors are small compared to the molars, but the canines are relatively large, with sexual dimorphism in some species. Cutting crests on the molars are well developed in some species, and the two halves of the mandible were fused at the midline in some species. Some adapoids were quite small (<em>Anchomomys <\/em>at a little over 100 g), and some were quite large (<em>Magnadapis<\/em> at 10 kg; 22 lbs.). Furthermore, the spaces and attachment features for the chewing muscles were truly enormous in some species, suggesting that these muscles were very large and powerful. Taken together, this suggests an overall adaptive profile of diurnal herbivory. The canine sexual dimorphism in some species suggests a possible mating pattern of polygyny, as males in polygynous primate species often compete with each other for mates and have especially large canine teeth.<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"548\"]<img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image18-1.jpg\" alt=\"Three partial animal crania.\" width=\"548\" height=\"350\" \/> Figure 9.9: Representative crania of Adapidae from Museum d\u2019Histoire Naturelle Victor Brun, a natural history museum in Montauban, France. The white scale bar is 1 cm long. Credit: <a class=\"rId43\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Representative crania of adapids (European adapoids, (Figure 8.7)<\/a> from the <a class=\"rId44\" href=\"https:\/\/www.museum.montauban.com\/\">Museum d\u2019Histoire Naturelle Victor Brun in Montauban, France<\/a> original to <a class=\"rId45\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology <\/a>by Jonathan M. G. Perry is under a <a class=\"rId46\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.[\/caption]\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"547\"]<img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image19-2.jpg\" alt=\"Side views of small rodentlike skeleton with long tail.\" width=\"547\" height=\"525\" \/> Figure 9.10: Darwinius masillae, a member of the Caenopithecidae. The slab on the left is Plate A and the slab on the right is Plate B. The parts of the skeleton in B that are outside of the dashed lines were fabricated. Credit: <a class=\"rId48\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Darwinius%20masillae%20holotype%20slabs.jpg\">Darwinius masillae holotype slabs<\/a> by Jens L. Franzen, Philip D. Gingerich, J\u00f6rg Habersetzer1, J\u00f8rn H. Hurum, Wighart von Koenigswald, B. Holly Smith is under a <a class=\"rId49\" href=\"https:\/\/creativecommons.org\/licenses\/by\/2.5\/legalcode\">CC BY 2.5 License<\/a>. Originally from Franzen Et al. 2009.[\/caption]\r\n<h4 class=\"import-Normal\"><em>Omomyoid Diversity<\/em><\/h4>\r\n<p class=\"import-Normal\">Like adapoids, omomyoids appeared suddenly at the start of the Eocene and then became very diverse with most species dying out before the Oligocene. Omomyoids are known from thousands of jaws with teeth, relatively complete skulls for about a half-dozen species, and very little postcranial material. Omomyoids were relatively small primates, with the largest being less than three kilograms (approximately 7 lbs.; <em>Macrotarsius montanus<\/em>). All known crania possess a postorbital bar, which in some has been described as \u201cincipient closure.\u201d Some\u2014but not all\u2014known crania have an elongated bony ear tube extending lateral to the location of the eardrum, a feature seen in living tarsiers and <strong>[pb_glossary id=\"2568\"]catarrhines[\/pb_glossary]<\/strong>. The anterior teeth tend to be large, with canines that are usually not much larger than the incisors. Often it is difficult to distinguish closely related species using molar morphology, but the premolars tend to be distinct from one species to another. The postcranial skeleton of most omomyoids shows hallmarks of leaping behaviour reminiscent of that of tarsiers. In North America, omomyoids became very diverse and abundant. In fact, omomyoids from Wyoming are sufficiently abundant and from such stratigraphically controlled conditions that they have served as strong evidence for the gradual evolution of anatomical traits over time (Rose and Bown 1984).<\/p>\r\n<p class=\"import-Normal\"><em>Teilhardina <\/em>(Figure 9.11; see Figure 9.2) is one of the earliest and arguably the most plesiomorphic of omomyoids. <em>Teilhardina<\/em> has several species, most of which are from North America, with one from Europe (<em>T. belgica<\/em>) and one from Asia (<em>T. asiatica<\/em>). The species of this genus are anatomically similar and the deposits from which they are derived are roughly contemporaneous. Thus, this small primate likely dispersed across the northern continents very rapidly (Smith Et al. 2006).<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"545\"]<img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image3-1.jpg\" alt=\"World map with primates jumping across forested areas.\" width=\"545\" height=\"289\" \/> Figure 9.11: A map of the world during the early Eocene showing one hypothesis for the direction of dispersal of the omomyoid Teilhardina. The map depicts primates hopping from continent to continent (East to West) via the forest corridors at high latitudes. Credit: <a href=\"https:\/\/www.pnas.org\/content\/103\/30\/11223\">Paleogeographic map showing hypothetical migration routes of Teilhardina (Figure 1)<\/a> by Thierry Smith, Kenneth D. Rose, and Philip D. Gingerich. 2006. <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">Proceedings of the National Academy of Sciences of the United States of America <\/a>103 (30): 11223\u201311227. Copyright (2006) National Academy of Sciences. Image <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">is used for non-commercial and educational purposes as outlined by PNAS.<\/a>[\/caption]\r\n<h2 class=\"import-Normal\">The Emergence of Modern Primate Groups<\/h2>\r\n<h3 class=\"import-Normal\"><strong>Origins of Crown Strepsirrhines<\/strong><\/h3>\r\n<p class=\"import-Normal\">Until the turn of this century, very little was known about the origins of the <strong>[pb_glossary id=\"1698\"]crown[\/pb_glossary]<\/strong> (living) strepsirrhines. The Quaternary record of Madagascar contains many amazing forms of lemurs, including giant sloth-like lemurs, lemurs with perhaps monkey-like habits, lemurs with koala-like habits, and even a giant aye-aye (Godfrey and Jungers 2002). However, in Madagascar, early Tertiary continental sediments are lacking, and there is no record of lemur fossils before the Pleistocene.<\/p>\r\n<p class=\"import-Normal\">The fossil record of galagos is slightly more informative. Namely, there are Miocene African fossils that are very likely progenitors of lorisids (Simpson 1967). However, these are much like modern galagos and do not reveal anything about the relationship between crown strepsirrhines and Eocene fossil primates (but see below regarding <em>Propotto<\/em>). A similar situation exists for lorises in Asia: there are Miocene representatives, but these are substantially like modern lorises. The discovery of the first definite [pb_glossary id=\"1699\"]<strong>toothcomb<\/strong>[\/pb_glossary] canine (a hallmark of stresirrhines) in 2003 provided the \u201csmoking gun\u201d for the origin of crown strepsirrhines (Seiffert Et al. 2003). Recently, several other African primates have been recognized as having strepsirrhine affinities (Marivaux Et al. 2013; Seiffert 2012). The enigmatic Fayum primate <em>Plesiopithecus<\/em> is known from a skull that has been compared to aye-ayes and to lorises (Godinot 2006; Simons and Rasmussen 1994a).<\/p>\r\n<p class=\"import-Normal\">The now-recognized diversity of stem strepsirrhines from the Eocene and Oligocene of Afro-Arabia is strong evidence to suggest that strepsirrhines originated in that geographic area. This implies that lorises dispersed to Asia subsequent to an African origin. It is unknown what the first strepsirrhines in Madagascar were like. However, it seems likely that the lemuriform-lorisiform split occurred in continental Africa, followed by dispersal of lemuriform stock to Madagascar. Recent evidence suggests that <em>Propotto<\/em>, a Miocene primate from Kenya originally described as a potto antecedent, actually forms a clade with <em>Plesiopithecus <\/em>and the aye-aye; this might suggest that strepsirrhines dispersed to Madagascar from continental Africa more than once (Gunnell Et al. 2018).<\/p>\r\n\r\n<h3 class=\"import-Normal\"><strong>The Fossil Record of Tarsiers<\/strong><\/h3>\r\n<p class=\"import-Normal\">Tarsiers are so unusual that they fuel major debates about primate taxonomy. Tarsiers today are moderately diverse but geographically limited and not very different in their ecological habits\u2014especially considering that the split between them and their nearest living relative probably occurred over 50 million years ago. If omomyoids are excluded, then the fossil record of tarsiers is very limited. Two fossil species from the Miocene of Thailand have been placed in the genus <em>Tarsius<\/em>, as has an Eocene fossil from China (Beard Et al. 1994). These, and <em>Xanthorhysis<\/em> from the Eocene of China, are all very tarsier-like. In fact, it is striking that <em>Tarsius eocaenus<\/em> from China was already so tarsier-like as early as the Eocene. This suggests that tarsiers achieved their current morphology very early in their evolution and have remained more or less the same while other primates changed dramatically. Two additional genera, <em>Afrotarsius<\/em> from the Oligocene of Egypt and Libya and <em>Afrasia<\/em> from the Eocene of Myanmar, have also been implicated in tarsier origins, though the relationship between them and tarsiers is unclear (Chaimanee Et al. 2012). More recently, a partial skeleton of a small Eocene primate from China, <em>Archicebus achilles<\/em> (dated to approximately 55.8 million to 54.8 million years ago), was described as the most basal tarsiiform (Ni Et al. 2013). This primate is reconstructed as a diurnal insectivore and an arboreal quadruped that did some leaping\u2014but not to the specialized degree seen in living tarsiers. The anatomy of the eye in living tarsiers suggests that their lineage passed through a diurnal stage, so <em>Archicebus<\/em> (and diurnal omomyoids) might represent such a stage.<\/p>\r\n\r\n<h3 class=\"import-Normal\"><strong>Climate Change and the Paleogeography of Modern Primate Origins<\/strong><\/h3>\r\n<p class=\"import-Normal\">Changing global climate has had profound effects on primate dispersal patterns and ecological habits over evolutionary time. Primates today are strongly tied to patches of trees and particular plant parts such as fruits, seeds, and immature leaves. It is no surprise, then, that the distribution of primates mirrors the distribution of forests. Today, primates are most diverse in the tropics, especially in tropical rainforests. Global temperature trends across the Tertiary have affected primate ranges. Following the Cretaceous-Tertiary extinction event, cooler temperatures and greater seasonality characterized the Paleocene. In the Eocene, temperatures (and probably rainfall) increased globally and rainforests likely extended to very high latitudes. During this time, euprimates became diverse. With cooling and increased aridity at the end of the Eocene, many primate extinctions occurred in the northern continents and the surviving primates were confined to lower latitudes in South America, Afro-Arabia, Asia, and southern Europe. Among these survivors are the progenitors of the living groups of primates: lemurs and lorises, tarsiers, [pb_glossary id=\"1700\"]<strong>platyrrhines<\/strong>[\/pb_glossary] (monkeys of the Americas), and catarrhines (monkeys and apes of Africa and Asia) (Figure 9.12).<\/p>\r\n\r\n\r\n[caption id=\"attachment_277\" align=\"aligncenter\" width=\"539\"]<img class=\"wp-image-264\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image2-5-e1691791570984.png\" alt=\"Map of world with gray continents.\" width=\"539\" height=\"306\" \/> Figure 9.12: Map of key localities of early anthropoids on land that becomes Africa and southern Asia. Credit: <a class=\"rId56\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Oligocene Map with Key Early Anthropoid Localities (Figure 8.10)<\/a> original to <a class=\"rId57\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Elyssa Ebding at <a class=\"rId58\" href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\">GeoPlace, California State University, Chico<\/a> is under a <a class=\"rId59\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Localities based on Fleagle 2013, 265.[\/caption]\r\n<h3 class=\"import-Normal\"><strong>Competing Hypotheses for the Origin of Anthropoids<\/strong><\/h3>\r\n<p class=\"import-Normal\">There is considerable debate among paleoanthropologists as to the geographic origins of anthropoids. In addition, there is debate regarding the source group for anthropoids. Three different hypotheses have been articulated in the literature. These are the adapoid origin hypothesis, the omomyoid origin hypothesis, and the tarsier origin hypothesis (Figure 9.13).<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"419\"]<img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image24-1-1.jpg\" alt=\"Diagrams show three relationships among primate groups.\" width=\"419\" height=\"742\" \/> Figure 9.13: Competing models of anthropoid origins. Branch lengths are not to scale. The omomyoid origin model and tarsier origin model do not make specific reference to the evolutionary position of strepsirrhines; however, they were included here for completeness. <a href=\"https:\/\/docs.google.com\/document\/d\/1VUDKMBJYS_jNONjLxT04jQN0_z9Ua50BRN6auGSHUuU\/edit\">A full text description of this image is available<\/a>. Credit: <a class=\"rId61\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Competing Trees for Anthropoid Origins (Figure 8.11)<\/a> original to <a class=\"rId62\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Jonathan M. G. Perry is under a <a class=\"rId63\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.[\/caption]\r\n<h4 class=\"import-Normal\"><em>Adapoid Origin Hypothesis<\/em><\/h4>\r\n<p class=\"import-Normal\">Resemblances between some adapoids and some extant anthropoids include fusion of the [pb_glossary id=\"1702\"]<strong>mandibular symphysis<\/strong>[\/pb_glossary], overall robusticity of the chewing system, overall large body size, features that signal a diurnal lifestyle (like relatively small eye sockets), and ankle bone morphology. Another feature in common is canine sexual dimorphism, which is present in some species of adapoids (probably) and in several species of anthropoids.<\/p>\r\n<p class=\"import-Normal\">These features led some paleoanthropologists in the last half of the 20th century to suggest that anthropoids came from adapoid stock (Gingerich 1980; Simons and Rasmussen 1994b). One of the earliest supporters of the link between adapoids and anthropoids was Hans Georg Stehlin, who described much of the best material of adapoids and compared these Eocene primates to South American monkeys (Stehlin 1912). In more recent times, the adapoid origin hypothesis was reinforced by resemblances between these European adapoids (especially <em>Adapis <\/em>and <em>Leptadapis<\/em>) and some early anthropoids from the Fayum Basin (e.g., <em>Aegyptopithecus<\/em>, see below; Figure 9.14).<\/p>\r\n\r\n<div style=\"text-align: left\">\r\n<table class=\"aligncenter\" style=\"width: 473.25pt\"><caption>Figure 9.14: Families of early anthropoids with example genera and traits: a table. Credit: Early anthropoids table original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Jonathan M. G. Perry and Stephanie L. Canington is under a <a class=\"rId64\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Content derived from Fleagle 2013.<\/caption>\r\n<thead>\r\n<tr style=\"height: 25pt\">\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Family<\/strong><\/p>\r\n&nbsp;<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Genera<\/strong><\/p>\r\n&nbsp;<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Morphology<\/strong><\/p>\r\n&nbsp;<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Location<\/strong><\/p>\r\n&nbsp;<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Age<\/strong><sup><strong>1<\/strong><\/sup><\/p>\r\n&nbsp;<\/td>\r\n<\/tr>\r\n<\/thead>\r\n<tbody>\r\n<tr class=\"Table3-R\" style=\"height: 18pt\">\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Propliopithecidae<sup>2<\/sup><\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\"><em>Aegyptopithecus<\/em><\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Large size. Cranial sexual dimorphism, large canines. Robust jaws and rounded molars. Partially ossified ear tube (in some). Robust skeleton; quadruped.<\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Africa<\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Late Eocene to Early Oligocene<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table3-R\" style=\"height: 16pt\">\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Parapithecidae<sup>3<\/sup><\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\"><em>Apidium<\/em><\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Medium size. Retention of three premolars per quadrant. Rounded molars and premolars with large central cusps. Adaptations for leaping in the lower limb.<\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Africa<\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Late Eocene to Late Oligocene<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table3-R\" style=\"height: 16pt\">\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Proteopithecidae<sup>4<\/sup><\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\"><em>Proteopithecus<\/em><\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Small size. Retention of three premolars per quadrant. Arboreal quadrupeds that ate fruit.<\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Africa<\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Late Eocene<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table3-R\" style=\"height: 16pt\">\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Oligopithecidae<sup>5<\/sup><\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\"><em>Catopithecus<\/em><\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Small size. Skull has postorbital septum and unfused mandible. Deep jaws. Diet of fruits. Generalized quadruped.<\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Africa<\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Late Eocene<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table3-R\" style=\"height: 16pt\">\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Eosimiidae<\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\"><em>Eosimias<\/em><\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Deep jaw with vertical unfused symphysis. Pointed incisors and canines. Crowded premolars.<\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Asia<\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Middle Eocene<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table3-R\" style=\"height: 16pt\">\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Amphipithecidae<sup>6<\/sup><\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\"><em>Pondaungia<\/em><\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Deep jaws. Molars generally rounded with wide basins.<\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Asia<\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\r\n<p class=\"import-Normal\">Middle Eocene to Early Oligocene<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table3-R\" style=\"height: 1pt\">\r\n<td class=\"Table3-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\" colspan=\"4\">\r\n<p class=\"import-Normal\"><sup>1<\/sup> Derived from Fleagle 2013.<\/p>\r\n<p class=\"import-Normal\"><sup>2<\/sup> See Gebo and Simons 1987 and Simons Et al. 2007.<\/p>\r\n<p class=\"import-Normal\"><sup>3<\/sup> See Feagle and Simons 1995 and Simons 2001.<\/p>\r\n<p class=\"import-Normal\"><sup>4<\/sup> See Simons and Seiffert 1999.<\/p>\r\n<p class=\"import-Normal\"><sup>5<\/sup> See Simons and Rasmussen 1996.<\/p>\r\n<p class=\"import-Normal\"><sup>6<\/sup> See Kay Et al. 2004.<\/p>\r\n<\/td>\r\n<td class=\"Table3-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\">\r\n<p class=\"import-Normal\"><\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr>\r\n<td><\/td>\r\n<td><\/td>\r\n<td><\/td>\r\n<td><\/td>\r\n<td><\/td>\r\n<\/tr>\r\n<\/tbody>\r\n<\/table>\r\n<\/div>\r\n<p class=\"import-Normal\">Unfortunately for the adapoid hypothesis, most of the shared features listed above probably emerged independently in the two groups as adaptations to a diet of hard and\/or tough foods. For example, fusion of the mandibular symphysis likely evolved as a means to strengthen the jaw against forces that would pull the two halves away from each other, in the context of active chewing muscles on both sides of the head generating great bite forces. This context would also favour the development of robust jaws, large chewing muscles, shorter faces, and some other features shared by some adapoids and some anthropoids.<\/p>\r\n<p class=\"import-Normal\">As older and more plesiomorphic anthropoids were found in the Fayum Basin, it became clear that the earliest anthropoids from Africa did not possess these features of jaw robusticity (Seiffert Et al. 2009). Furthermore, many adapoids never evolved these features. Fusion of the mandibular symphysis in adapoids is actually quite different from that in anthropoids and probably occurred during juvenile development in the former (Beecher 1983; Ravosa 1996). Eventually, the adapoid origin hypothesis fell out of favour among most paleoanthropologists, although the description of <em>Darwinius<\/em> is a recent revival of that idea (Franzen Et al. 2009; but see Seiffert Et al. 2009, Williams Et al. 2010b).<\/p>\r\n\r\n<h4 class=\"import-Normal\"><em>Omomyoid Origin Hypothesis<\/em><\/h4>\r\n<p class=\"import-Normal\">Similarities in cranial and hindlimb morphology between some omomyoids and extant tarsiers have led to the suggestion that tarsiers arose from some kind of omomyoid. In particular, <em>Necrolemur<\/em> has many features in common with tarsiers, as does the North American <em>Shoshonius<\/em>, which is known from a few beautifully preserved (although distorted) crania. Tarsiers and <em>Shoshonius <\/em>share exclusively some features of the base of the cranium; however, <em>Shoshonius<\/em> does not have any sign of postorbital closure, and it lacks the bony ear tube of tarsiers. Nevertheless, some of the resemblances between some omomyoids and tarsiers suggest that tarsiers might have originated from within the Omomyoidea (Beard 2002; Beard and MacPhee 1994). In this scenario, although living tarsiers and living anthropoids might be sister taxa, they might have evolved from different omomyoids, possibly separated from each other by more than 50 million years of evolution, or from anthropoids evolved from some non-omomyoid fossil group. The arguments against the omomyoid origin hypothesis are essentially the arguments <em>for<\/em> the tarsier origin hypothesis (see below). Namely, tarsiers and anthropoids share many features (especially of the soft tissues) that must have been retained for many millions of years or must have evolved convergently in the two groups. Furthermore, a key hard-tissue feature shared between the two extant groups, the postorbital septum, was not present in any omomyoid. Therefore, that feature must have arisen convergently in the two extant groups or must have been lost in omomyoids. Neither scenario is very appealing, although recent arguments for <strong>convergent evolution<\/strong> of the postorbital septum in tarsiers and anthropoids have arisen from embryology and histology of the structure (DeLeon Et al. 2016).<\/p>\r\n\r\n<h4 class=\"import-Normal\"><em>Tarsier Origin Hypothesis<\/em><\/h4>\r\n<p class=\"import-Normal\">Several paleoanthropologists have suggested that there is a relationship between tarsiers and anthropoids to the exclusion of omomyoids and adapoids (e.g., Cartmill and Kay 1978; Ross 2000; Williams and Kay 1995). Tarsiers and anthropoids today share several traits, including many soft-tissue features related to the olfactory system (e.g., the loss of a hairless external nose and loss of the median cleft running from the nose to the mouth, as possessed by strepsirrhines), and aspects of the visual system (e.g., the loss of a reflective layer at the back of the eye, similarities in carotid circulation to the brain, and mode of placentation). Unfortunately, none of these can be assessed directly in fossils. Some bony similarities between tarsiers and anthropoids include an extra air-filled chamber below the middle ear cavity, reduced bones within the nasal cavity, and substantial postorbital closure; these can be assessed in fossils, but the distribution of these traits in omomyoids does not yield clear answers. Furthermore, several similarities between tarsiers and anthropoids are probably due to similarities in sensory systems, which might have evolved in parallel for ecological reasons. Although early attempts to resolve the crown primates with molecular data were sometimes equivocal or in disagreement with one another, more recent analyses (including those of short interspersed elements) suggest that tarsiers and anthropoids are sister groups to the exclusion of lemurs and lorises (Williams Et al. 2010a). However, this does not address omomyoids, all of which are far too ancient for DNA extraction.<\/p>\r\n<p class=\"import-Normal\">The above three hypotheses are not the only possibilities for anthropoid origins. It may be that anthropoids are neither the closest sister group of tarsiers, nor evolved from adapoids or omomyoids. In recent years, two new groups of Eocene Asian primates have been implicated in the origin of anthropoids: the eosimiids and the amphipithecids. It is possible that one or the other of these two groups gave rise to anthropoids. Regardless of the true configuration of the tree for crown primates, the three major extant groups probably diverged from each other quite long ago (Seiffert Et al. 2004).<\/p>\r\n\r\n<h3 class=\"import-Normal\"><strong>Early Anthropoid Fossils in Africa<\/strong><\/h3>\r\n[caption id=\"\" align=\"aligncenter\" width=\"526\"]<img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image7-2.jpg\" alt=\"People digging in a sandy desert.\" width=\"526\" height=\"352\" \/> Figure 9.15: Egyptian workers sweeping Quarry I in the Fayum Basin (2004). This technique, called wind harvesting, removes the desert crust and permits wind to blow out fine sediment and reveal fossils. Credit: <a class=\"rId66\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Egyptian workers sweeping Quarry I in the Fayum Basin (2004, Figure 8.12)<\/a> by Jonathan M. G. Perry is under a <a class=\"rId67\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.[\/caption]\r\n\r\n[caption id=\"\" align=\"alignleft\" width=\"280\"]<img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image14-2.jpg\" alt=\"A person using a tool to expose bone in sand.\" width=\"280\" height=\"423\" \/> Figure 9.16: Elwyn Laverne Simons excavating Aegyptopithecus in the Fayum Basin. Credit: <a class=\"rId69\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Elwyn Laverne Simons in the Fayum Basin (Figure 8.13)<\/a> used by permission of the <a class=\"rId70\" href=\"https:\/\/lemur.duke.edu\/\">Duke Lemur Center,<\/a> Division of Fossil Primates, is under a <a class=\"rId71\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.[\/caption]\r\n<p class=\"import-Normal\">The classic localities yielding the greatest wealth of early anthropoid fossils are those from the Fayum Basin in Egypt (Simons 2008; Figure 9.15). The Fayum is a veritable oasis of fossil primates in an otherwise spotty early Tertiary African record. Since the 1960s, teams led by E. L. Simons have discovered several new species of early anthropoids, some of which are known from many parts of the skeleton and several individuals (Figure 9.16).<\/p>\r\n<p class=\"import-Normal\">The Fayum Jebel Qatrani Formation and Birket Qarun Formation between them have yielded a remarkable array of terrestrial, arboreal, and aquatic mammals. These include ungulates, bats, sea cows, elephants, hyraces, rodents, whales, and primates. Also, many other vertebrates, like water birds, were present. The area at the time of deposition (Late Eocene through Early Oligocene) was probably very wet, with slow-moving rivers, standing water, swampy conditions, and lots of trees (see Bown and Kraus 1988). In short, it was an excellent place for primates.<\/p>\r\n\r\n<h4 class=\"import-Normal\"><em>General Morphology of Anthropoids<\/em><\/h4>\r\n<p class=\"import-Normal\">The anthropoids known from the Fayum (and their close relatives from elsewhere in East Africa and Afro-Arabia) bear many of the anatomical hallmarks of extant anthropoids; however, there are plesiomorphic forms in several families that lack one or more anthropoid traits. All Fayum anthropoids known from skulls possess postorbital closure, most had fused mandibular symphyses, and most had ring-like [pb_glossary id=\"1704\"]<strong>ectotympanic<\/strong> [\/pb_glossary] bones. Tooth formulae were generally either 2.1.3.3 or 2.1.2.3. Fayum anthropoids ranged in size from the very small <em>Qatrania<\/em> and <em>Biretia <\/em>(less than 500 g) to the much-larger <em>Aegyptopithecus<\/em> (approximately 7 kg; 15 lbs.). Fruit was probably the main component of the diet for most or all of the anthropoids, with some of them supplementing with leaves (Kay and Simons 1980; Kirk and Simons 2001; Teaford Et al. 1996). Most Fayum anthropoids were probably diurnal above-branch quadrupeds. Some of them (e.g., <em>Apidium<\/em>; see Figure 9.14) were probably very good leapers (Gebo and Simons 1987), but none show specializations for gibbon-style suspensory locomotion. Some of the Fayum anthropoids are known from hundreds of individuals, permitting the assessment of individual variation, sexual dimorphism, and in some cases growth and development. The description that follows provides greater detail for the two best known Fayum anthropoid families, the Propliopithecidae and the Parapithecidae; the additional families are summarized briefly.<\/p>\r\n\r\n<h4 class=\"import-Normal\"><em>Fayum Anthropoid Families<\/em><\/h4>\r\n<p class=\"import-Normal\">The Propliopithecidae (see Figure 9.14) include the largest anthropoids from the fauna, and they are known from several crania and some postcranial elements. They have been suggested to be stem catarrhines, although perhaps near the split between catarrhines and platyrrhines. The best known propliopithecid is <em>Aegyptopithecus<\/em>, known from many teeth, crania, and postcranial elements (Figure 9.17) .<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"alignright\" width=\"431\"]<img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-2-1.jpg\" alt=\"Two animal skull side views.\" width=\"431\" height=\"281\" \/> Figure 9.17: Female (left) and male (right) skull material for Aegyptopithecus zeuxis. The mandibles are not associated with the crania. Credit: <a href=\"https:\/\/www.pnas.org\/doi\/full\/10.1073\/pnas.0703129104#supplementary-materials\">Female and male cranium of A. zeuxi (03129Fig5, Supporting Information)<\/a> by Elwyn L. Simons, Erik R. Seiffert, Timothy M. Ryan, and Yousry Attia. 2007. <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">Proceedings of the National Academy of Sciences of the United States of America<\/a> 104 (21): 8731\u20138736. Copyright (2007) National Academy of Sciences. Image <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">is used for non-commercial and educational purposes as outlined by PNAS.<\/a>[\/caption]\r\n<p class=\"import-Normal\">Parapithecidae are an extremely abundant and unusual family of anthropoids from the Fayum. The parapithecid <em>Apidium<\/em> is known from many jaws with teeth, crushed and distorted crania, and several skeletal elements. <em>Parapithecus<\/em> is known from cranial material including a beautiful, undistorted cranium. This genus shows extreme reduction of the incisors, including complete absence of the lower incisors in <em>P. grangeri <\/em>(Simons 2001). This trait is unique among primates. Parapithecids were once thought to be the ancestral stock of platyrrhines; however, their platyrrhine-like features are probably ancestral retentions, so the most conservative approach is to consider them stem anthropoids.<\/p>\r\n<p class=\"import-Normal\">The Proteopithecidae were small frugivores that probably mainly walked along horizontal branches on all fours. They are considered stem anthropoids. The best known genus, <em>Proteopithecus<\/em>, is represented by dentitions, crania, and post-cranial elements.<\/p>\r\n<p class=\"import-Normal\">The Oligopithecidae share a mixture of traits that makes them difficult to classify more specifically within anthropoids. The best known member, <em>Catopithecus<\/em>, is known from crania that demonstrate a postorbital septum and from mandibles that lack symphyseal fusion. They share the catarrhine tooth formula of 2.1.2.3 and have a canine honing complex that involves the anterior lower premolar. The postcranial elements known for the group suggest generalized arboreal quadrupedalism. The best known member, <em>Catopithecus<\/em>, is known from crania that demonstrate a postorbital septum and from mandibles that lack symphyseal fusion (Simons and Rasmussen 1996). The jaws are deep, with broad muscle attachment areas and crested teeth. <em>Catopithecus<\/em> was probably a little less than a kilogram in weight.<\/p>\r\n<p class=\"import-Normal\">Other genera of putative anthropoids from the Fayum include the very poorly known <em>Arsinoea<\/em>, the contentious <em>Afrotarsius<\/em>, and the enigmatic <em>Nosmips<\/em>. The last of these possesses traits of several major primate <strong>[pb_glossary id=\"1705\"]clades[\/pb_glossary]<\/strong> and defies classification (Seiffert Et al. 2010).<\/p>\r\n\r\n<h3 class=\"import-Normal\"><strong>Early Anthropoid Fossils in Asia<br style=\"clear: both\" \/><\/strong><\/h3>\r\n<p class=\"import-Normal\">For the last half of the 1900s, researchers believed that Africa was the unquestioned homeland of early anthropoids (see Fleagle and Kay 1994). However, two very different groups of primates from Asia soon began to change that. One was an entirely new discovery (Eosimiidae), and the other was a poorly known group discovered decades prior (Amphipithecidae). Soon, attention on anthropoid origins began to shift eastward (see Ross and Kay 2004; Simons 2004). If anthropoids arose in Asia instead of Africa, then this implies that the African early anthropoids either emigrated from Asia or evolved their anthropoid traits in parallel with living anthropoids.<\/p>\r\n\r\n<h4 class=\"import-Normal\"><em>Eosimiids<\/em><\/h4>\r\n<p class=\"import-Normal\">First described in the 1990s, the eosimiids are best represented by <em>Eosimias <\/em>(see Figure 9.14; Figure 9.18). This tiny \u201cdawn monkey\u201d is known from relatively complete jaws with teeth, a few small fragments of the face, and some postcranial elements (Beard Et al. 1994; Beard Et al. 1996; Gebo Et al. 2000). <em>Eosimias<\/em> (along with the other less-well-known genera in its family) bears some resemblance to tarsiers as well as anthropoids. Unfortunately, no good crania are known for this family, and the anatomy of, for example, the posterior orbital margin could be very revealing as to higher-level relationships.<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"550\"]<img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image16-1-1.jpg\" alt=\"Red-colored lower jaw of an animal.\" width=\"550\" height=\"232\" \/> Figure 9.18: Cast of the right half of the mandible of Eosimias centennicus, type specimen. The white scale bar is 1 cm long. Credit: <a class=\"rId74\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Cast of the right half of the mandible of <\/a><a class=\"rId75\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\"><em>Eosimias centennicus <\/em><\/a><a class=\"rId76\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">(Figure 8.15),<\/a> type specimen, from K. D. Rose cast collection, photo by Jonathan M. G. Perry is under a <a class=\"rId77\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.[\/caption]\r\n<h4 class=\"import-Normal\"><em>Amphipithecids<\/em><\/h4>\r\n<p class=\"import-Normal\">Amphipithecids are small- to medium-size primates (up to 10 kg; 22 lbs.). Most are from the Eocene Pondaung Formation in Myanmar (Early\u2013Middle Eocene), but one genus is known from Thailand. Some dental similarities with anthropoids were noted early on, such as deep jaws and wide basins that separate low molar cusps. The best known genera were <em>Pondaungia<\/em> and <em>Amphipithecus <\/em>(Ciochon and Gunnell 2002; see Figure 9.14). Another amphipithecid, <em>Siamopithecus<\/em> from Thailand, has very rounded molars and was probably a seed-eater (Figure 9.19). In addition to teeth and jaws, some cranial fragments, ankle material, and ends of post-cranial bones have been found for <em>Pondaungia<\/em>. There are important resemblances between these bones of <em>Pondaungia<\/em> and those of adapoids, suggesting adapoid affinities for the amphipithecidae. This would imply that the resemblances with anthropoids in the teeth are convergent, based on similarities in diet (see Ciochon and Gunnell 2002). Unfortunately, the association between post-cranial bones and teeth is not definite. With other primates in these faunas (including eosimiids), one cannot be certain that the post-cranial bones belong with the teeth. Some researchers suggest that some bones belong to a sivaladapid (or asiadapid) and others to an early anthropoid (Beard Et al. 2007; Marivaux Et al. 2003). Additional well-associated material of amphipithecids would help to clear up this uncertainty.<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"505\"]<img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image15-2.jpg\" alt=\"Four casts of jawbone fragments with teeth.\" width=\"505\" height=\"368\" \/> Figure 9.19: Casts of representative amphipithecid material. A. Pondaungia cotteri right lower jaw fragment with m2 and m3. B. Siamopithecus eocaenus right upper jaw fragment with p4-m3. C. S. eocaenus right lower jaw fragment with partial m1, m2, and m3 in lateral view. D. Same as in C but occlusal view. White scale bars are 1 cm long. Credit: <a class=\"rId79\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Casts of representative amphipithecid material (Figure 8.16)l<\/a> from K. D. Rose cast collection, photo by Jonathan M. G. Perry is under a <a class=\"rId80\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.[\/caption]\r\n<h3 class=\"import-Normal\"><strong>Platyrrhine Dispersal to South America<\/strong><\/h3>\r\n<p class=\"import-Normal\">Today there is an impressive diversity of primates in South and Central America. These are considered to be part of a single clade, the Platyrrhini. Primates colonized South America sometime in the Eocene from an African source. In the first half of the 20th century, the source of platyrrhines was a matter of major debate among paleontologists, with some favouring a North American origin (e.g., Simpson 1940).<\/p>\r\n<p class=\"import-Normal\">Part of the reason for this debate is that South America was an island in the Eocene. Primates needed to cross open ocean to get there from either North America or Africa, although the distance from the former was shorter. Morphology yields clues to platyrrhine origins. The first known primates in South America have more in common morphologically with African primates than with North American ones. At the time, anthropoids were popping up in North Africa, whereas the only euprimates in North America were adapoids and omomyoids. Despite lacking a bony ear tube, early platyrrhines shared a great deal with other anthropoids, including full postorbital closure and fusion of the mandibular symphysis.<\/p>\r\n<p class=\"import-Normal\">The means by which a population of small North African primates managed to disperse across the Atlantic and survive to colonize South America remains a mystery. The most plausible scenario is one of rafting. That is, primates must have been trapped on vegetation that was blown out to sea by a storm. The vegetation then became a sort of life raft, which eventually landed ashore, dumping its passengers in South America. Rodents probably arrived in South America in the same way (Antoine Et al. 2012).<\/p>\r\n<p class=\"import-Normal\">Once ashore, platyrrhines must have crossed South America fairly rapidly because the earliest-known primates from that continent are from Peru (Bond Et al. 2015). Soon after that, platyrrhines were in Bolivia, namely <em>Branisella<\/em>. By the Miocene, platyrrhines were living in extreme southern Argentina and were exploiting a variety of feeding niches. The Early Miocene platyrrhines were all somewhat plesiomorphic in their morphology, but some features that likely arose by ecological convergence suggest (to some) relationships with extant platyrrhine families. This has led to a lively debate about the pattern of primate evolution in South America (Kay 2015; Kay &amp; Fleagle 2010; Rosenberger 2010). By the Middle Miocene, clear representatives of modern families were present in a diverse fauna from La Venta, Colombia (Wheeler 2010). The Plio-Pleistocene saw the emergence of giant platyrrhines as well as several taxa of platyrrhines living on Caribbean islands (Cooke Et al. 2016).<\/p>\r\n<p class=\"import-Normal\">The story of platyrrhines seems to be one of amazing sweepstakes dispersal, followed by rapid diversification and widespread geographic colonization of much of South America. After that, dramatic extinctions resulted in the current, much-smaller geographic distribution of platyrrhines. These extinctions were probably caused by changing climates, leading to the contraction of forests. Platyrrhines dispersed to the Caribbean and to Central America, with subsequent extinctions in those regions that might have been related to interactions with humans. Unlike anthropoids of Africa and Asia, platyrrhines do not seem to have evolved any primarily terrestrial forms and so have always been highly dependent on forests.<\/p>\r\n\r\n<div class=\"textbox\">\r\n<h2 class=\"import-Normal\">Special Topic: Jonathan Perry and Primates of the Extreme South<\/h2>\r\n<p class=\"import-Normal\">Many primates are very vulnerable to ecological disturbance because they are heavily dependent on fruit to eat and trees to live in. This is one reason why so many primates are endangered today and why many of them went extinct due to climatic and vegetational changes in the past. I (Jonathan Perry) have conducted paleontological research focusing on primates that lived on the edge of their geographic distribution. This research has taken me to extreme environments in the Americas: southern Patagonia, the Canadian prairies, western Wyoming, and the badlands of eastern Oregon.<\/p>\r\n<p class=\"import-Normal\">Santa Cruz Province in Argentina is as far south as primates have ever lived. The Santa Cruz fauna of the Miocene has yielded a moderate diversity of platyrrhines, each with slightly different dietary adaptations. These include <em>Homunculus<\/em>, first described by Florentino Ameghino in 1891 (Figure 9.20). Recent fieldwork by my colleagues and I in Argentina has revealed several skulls of <em>Homunculus <\/em>as well as many parts of the skeleton (Kay Et al. 2012). The emerging profile of this extinct primate is one of a dedicated arboreal quadruped that fed on fruits and leaves. Many of the foods eaten by <em>Homunculus<\/em> must have been very tough and were probably covered and impregnated with grit; we suspect this because the cheek teeth are very worn down, even in young individuals, and because the molar tooth roots were very large, presumably to resist strong bite forces (Perry Et al. 2010, 2014).<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"497\"]<img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image9-2.jpg\" alt=\"An animal skull, a partial skull, and a fossil jaw with teeth.\" width=\"497\" height=\"634\" \/> Figure 9.20: Representative specimens of Homunculus patagonicus. A. Adult cranium in lateral view. B. Adult cranium surface reconstructed from microCT scans, with the teeth segmented out. C. Juvenile cranium. White scale bars are 1cm long. Credit: <a class=\"rId82\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Representative specimens of <\/a><a class=\"rId83\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\"><em>Homunculus patagonicus <\/em><\/a><a class=\"rId84\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">(Figure 8.17)<\/a> photo by Jonathan M. G. Perry is under a <a class=\"rId85\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.[\/caption]\r\n<p class=\"import-Normal\">I began working in Argentina while a graduate student at Duke University. I participated as a field assistant in a team led by my Ph.D. advisor, Richard F. Kay, and Argentine colleagues Sergio F. Vizca\u00edno and M. Susana Bargo. Most of the localities examined belong to a suite of beach sites known since the 1800s and visited by many field parties from various museums in the early 1900s. Since 2003, our international team of paleontologists from the U.S. and Argentina has visited these localities every single year (Figure 9.21). Over time, new fossils and new students have led to new projects and new approaches, including the use of microcomputed tomography (microCT) to visualize and analyse internal structures of the skeleton.<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"491\"]<img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image23.jpg\" alt=\"Sandy rocky coastline. People digging on a grassy hillside.\" width=\"491\" height=\"561\" \/> Figure 9.21: Field localities in Argentina and Canada. A. Ca\u00f1adon Palos locality, coastal Santa Cruz Province, Argentina. B. Swift Current Creek locality, southwest Saskatchewan, Canada. Credits: A. <a class=\"rId87\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Ca\u00f1adon Palos Field Locality in Argentina<\/a> by Jonathan M. G. Perry is under a <a class=\"rId88\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. B. <a class=\"rId89\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Swift Current Creek locality, Saskatchewan, Canada<\/a> by Jonathan M. G. Perry is under a <a class=\"rId90\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.[\/caption]\r\n\r\n<\/div>\r\n<h2 class=\"import-Normal\">Planet of Apes<\/h2>\r\n<h3 class=\"import-Normal\"><strong>Geologic Activity and Climate Change in the Miocene<\/strong><\/h3>\r\n<p class=\"import-Normal\">The Miocene Epoch was a time of mammalian diversification and extinction, global climate change, and ecological turnover. In the Miocene, there was an initial warming trend across the globe with the expansion of subtropical forests, followed by widespread cooling and drying with the retreat of tropical forests and replacement with more open woodlands and eventually grasslands. It was also a time of major geologic activity. On one side of the globe, South America experienced the rise of the Andes Mountains. On the other side, the Indian subcontinent collided with mainland Asia, resulting in the rise of the Himalayan Mountains. In Africa, volcanic activity promoted the development of the East African Rift System. Critical to the story of ape evolution was the exposure of an intercontinental landbridge between East Africa and Eurasia, permitting a true planet of apes (Figure 9.22).<\/p>\r\n\r\n\r\n[caption id=\"attachment_277\" align=\"aligncenter\" width=\"580\"]<img class=\"wp-image-273\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image21-3-e1691792198797.png\" alt=\"Map of world with gray continents.\" width=\"580\" height=\"335\" \/> Figure 9.22: Map of the world in the Miocene, highlighting fossil ape localities across Africa, southern Europe, and southern Asia. Credit: <a class=\"rId92\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Miocene Map with Fossil Ape Localities (Figure 8.19)<\/a> original to <a class=\"rId93\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Elyssa Ebding at <a class=\"rId94\" href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\">GeoPlace, California State University, Chico<\/a> is under a <a class=\"rId95\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Localities based on Fleagle 2013, 311.[\/caption]\r\n<h3 class=\"import-Normal\"><strong>Geographic Distribution: Africa, Asia, Europe<\/strong><\/h3>\r\n<p class=\"import-Normal\">The world of the Miocene had tremendous ape diversity compared to today. The earliest records of fossil apes are from Early Miocene deposits in Africa. However, something dramatic happened around 16 million years ago. With the closure of the ancient Tethys Sea, the subsequent exposure of the <em>Gomphotherium<\/em> Landbridge, and a period of global warming, the Middle\u2013Late Miocene saw waves of emigration of mammals (including primates) out of Africa and into Eurasia, with evidence of later African re-entry for some (Harrison 2010). Some of the mammals that dispersed from Africa to Eurasia and back were apes. Though most of these early apes left no modern descendants, some of them gave rise to the ancestors of modern apes\u2014including <strong>[pb_glossary id=\"800\"]hominins[\/pb_glossary]<\/strong> (Figure 9.23).<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"560\"]<img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image20-1.jpg\" alt=\"Miocene apes set against a geologic time scale.\" width=\"560\" height=\"796\" \/> Figure 9.23: Representative Miocene apes set against a geologic time scale. Credit: <a href=\"https:\/\/www.pnas.org\/content\/108\/14\/5554\">Range chart for Miocene hominoids of Western Eurasia (Figure 3)<\/a> by Isaac Casanovas-Vilar, David M. Alba, Miguel Garc\u00e9s, Josep M. Robles, and Salvador Moy\u00e0-Sol\u00e0. 2011. <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">Proceedings of the National Academy of Sciences of the United States of America<\/a> 108 (14): 5554-5559. Copyright (2011) National Academy of Sciences. Image <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">is used for non-commercial and educational purposes as outlined by PNAS.<\/a>[\/caption]\r\n<h3 class=\"import-Normal\"><strong>Where Are the Monkeys? Diversity in the Miocene<\/strong><\/h3>\r\n<p class=\"import-Normal\">Whereas the Oligocene deposits in the Fayum of Egypt have yielded the earliest-known catarrhine fossils, the Miocene demonstrates some diversification of Cercopithecoidea. However, compared to the numerous and diverse Miocene apes (see below), monkeys of the Miocene are very rare and restricted to a single extinct family, the Victoriapithecidae (Figure 9.24). This family contains the earliest definite cercopithecoids.<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"alignleft\" width=\"357\"]<img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image13-5.png\" alt=\"Front view of skull with pointed teeth.\" width=\"357\" height=\"322\" \/> Figure 9.25: Skull of Victoriapithecus macinnesi. Credit: Victoriapithecus macinnesi skull photo taken at the Musee d'Histoire Naturelle, Paris by Ghedoghedo is under a CC BY-SA 3.0 License.[\/caption]\r\n<p class=\"import-Normal\">These monkeys are found from northern and eastern Africa between 20 million and 12.5 million years ago (Miller Et al. 2009). The best known early African monkey is <em>Victoriapithecus <\/em>(Figure 9.25), a small-bodied (approximately 7 kg; 15 lbs.), small-brained monkey. <strong>[pb_glossary id=\"1708\"]Bilophodonty[\/pb_glossary]<\/strong>, known to be a hallmark of molar teeth of modern cercopithecoid, was present to some extent in Victoriapithecids. <em>Victoriapithecus<\/em> has been reconstructed as being more frugivorous and perhaps spent more time on the ground (terrestrial locomotion) than in the trees (arboreal locomotion; Blue Et al. 2006). The two major groups of cercopithecoids today are cercopithecines and colobines. The earliest records demonstrating clear members of each of these two groups are at the end of the Miocene. Examples include the early colobine <em>Microcolobus<\/em> from Kenya and the early cercopithecine <em>Pliopapio<\/em> from Ethiopia.<\/p>\r\n\r\n<div style=\"text-align: left\">\r\n<table class=\"aligncenter\" style=\"width: 473.25pt;height: 349px\"><caption>Figure 9.24: Some families of later anthropoids with example genera and traits: a table. Credit: Late anthropoids table original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Jonathan M. G. Perry and Stephanie L. Canington is under a <a class=\"rId100\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Content derived from Fleagle 2013.<\/caption>\r\n<thead>\r\n<tr style=\"height: 25pt\">\r\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 60px;width: 119.35px\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Family<\/strong><\/p>\r\n&nbsp;<\/td>\r\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 60px;width: 103.417px\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Genera<\/strong><\/p>\r\n&nbsp;<\/td>\r\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 60px;width: 191.65px\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Morphology<\/strong><\/p>\r\n&nbsp;<\/td>\r\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 60px;width: 67.3667px\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Location<\/strong><\/p>\r\n&nbsp;<\/td>\r\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 60px;width: 73.2167px\">\r\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Age<\/strong><sup><strong>1<\/strong><\/sup><\/p>\r\n&nbsp;<\/td>\r\n<\/tr>\r\n<\/thead>\r\n<tbody>\r\n<tr class=\"Table4-R\" style=\"height: 18pt\">\r\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 77px;width: 119.35px\">\r\n<p class=\"import-Normal\">Victoriapithecidae<sup>2<\/sup><\/p>\r\n<\/td>\r\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 77px;width: 103.417px\">\r\n<p class=\"import-Normal\"><em>Victoriapithecus<\/em><\/p>\r\n<\/td>\r\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 77px;width: 191.65px\">\r\n<p class=\"import-Normal\">Long, sloping face. Round, narrowly spaced orbits. Deep cheek bones. Well-developed sagittal crest.<\/p>\r\n<\/td>\r\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 77px;width: 67.3667px\">\r\n<p class=\"import-Normal\">Africa<\/p>\r\n<\/td>\r\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 77px;width: 73.2167px\">\r\n<p class=\"import-Normal\">Early to Middle Miocene<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table4-R\" style=\"height: 16pt\">\r\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 61px;width: 119.35px\">\r\n<p class=\"import-Normal\">Proconsulidae<sup>3<\/sup><\/p>\r\n<\/td>\r\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 61px;width: 103.417px\">\r\n<p class=\"import-Normal\"><em>Proconsul<\/em><\/p>\r\n<\/td>\r\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 61px;width: 191.65px\">\r\n<p class=\"import-Normal\">Short face. Generalized dentition. Arboreal quadruped. Probably tailless.<\/p>\r\n<\/td>\r\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 61px;width: 67.3667px\">\r\n<p class=\"import-Normal\">Africa and Arabia<\/p>\r\n<\/td>\r\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 61px;width: 73.2167px\">\r\n<p class=\"import-Normal\">Early to Middle Miocene<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table4-R\" style=\"height: 16pt\">\r\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 46px;width: 119.35px\">\r\n<p class=\"import-Normal\">Pongidae<\/p>\r\n<\/td>\r\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 46px;width: 103.417px\">\r\n<p class=\"import-Normal\"><em>Gigantopithecus<\/em><\/p>\r\n<\/td>\r\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 46px;width: 191.65px\">\r\n<p class=\"import-Normal\">Largest primate ever. Deep jaws and low rounded molars.<\/p>\r\n<\/td>\r\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 46px;width: 67.3667px\">\r\n<p class=\"import-Normal\">Asia<\/p>\r\n<\/td>\r\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 46px;width: 73.2167px\">\r\n<p class=\"import-Normal\">Miocene to Present<\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr class=\"Table4-R\" style=\"height: 1pt\">\r\n<td class=\"Table4-C\" style=\"border-color: #000000;border-style: solid none none;border-width: 0.5pt 0pt 0pt;padding: 0pt 5.4pt;height: 90px;width: 526.983px\" colspan=\"4\">\r\n<p class=\"import-Normal\"><sup>1<\/sup> Derived from Fleagle 2013.<\/p>\r\n<p class=\"import-Normal\"><sup>2<\/sup> See Benefit and McCrossin 1997 and Fleagle 2013.<\/p>\r\n<p class=\"import-Normal\"><sup>3<\/sup> See Begun 2007.<\/p>\r\n<\/td>\r\n<td class=\"Table4-C\" style=\"border-color: #000000;border-style: solid none none;border-width: 0.5pt 0pt 0pt;padding: 0pt 5.4pt;height: 90px;width: 73.2167px\">\r\n<p class=\"import-Normal\"><\/p>\r\n<\/td>\r\n<\/tr>\r\n<tr style=\"height: 15px\">\r\n<td style=\"height: 15px;width: 121.283px\"><\/td>\r\n<td style=\"height: 15px;width: 105.35px\"><\/td>\r\n<td style=\"height: 15px;width: 193.583px\"><\/td>\r\n<td style=\"height: 15px;width: 69.3px\"><\/td>\r\n<td style=\"height: 15px;width: 74.65px\"><\/td>\r\n<\/tr>\r\n<\/tbody>\r\n<\/table>\r\n<\/div>\r\n<h3><\/h3>\r\n<h3><\/h3>\r\n<h3><\/h3>\r\n<h3 class=\"import-Normal\"><strong>The Story of Us, the Apes<\/strong><\/h3>\r\n<h4 class=\"import-Normal\"><em>African Ape Diversity <\/em><\/h4>\r\n<p class=\"import-Normal\">The Early Miocene of Africa has yielded around 14 genera of early apes (Begun 2003). Many of these taxa have been reconstructed as frugivorous arboreal quadrupeds (Kay 1977). One of the best studied of these genera is the East African <em>Proconsul<\/em> (Family Proconsulidae; see Figure 9.24). Several species have been described, with body mass reconstructions ranging from 17 to 50 kg (approximately 37\u2013110 lbs.). A paleoenvironmental study reconstructed the habitat of <em>Proconsul <\/em>to be a dense, closed-canopy tropical forest (Michel Et al. 2014). No caudal vertebrae (tail bones) have been found in direct association with <em>Proconsul <\/em>post-crania, and the morphology of the sacrum is consistent with <em>Proconsul<\/em> lacking a tail (Russo 2016; Ward Et al. 1991).<\/p>\r\n<p class=\"import-Normal\">Overall, the African ape fossil record in the Late Miocene is sparse, with seven fossil localities dating between eleven and five million years ago (Pickford Et al. 2009). Nevertheless, most species of great apes live in Africa today. Where did the progenitors of modern African apes arise? Did they evolve in Africa or somewhere else? The paucity of apes in the Late Miocene of Africa stands in contrast to the situation in Eurasia. There, ape diversity was high. Furthermore, several Eurasian ape fossils show morphological affinities with modern hominoids (apes). Because of this, some paleoanthropologists suggest that the ancestors of modern African great apes recolonized Africa from Eurasia toward the end of the Miocene (Begun 2002). However, discoveries of Late Miocene hominoids like the Kenyan <em>Nakalipithecus<\/em> (9.9 million to 9.8 million years ago), the Ethiopian <em>Chororapithecus<\/em> (10.7 million to 10.1 million years ago), and the late-Middle Miocene Namibian <em>Otavipithecus<\/em> (13 million to 12 million years ago) fuel an alternative hypothesis\u2014namely that African hominoid diversity was maintained throughout the Miocene and that one of these taxa might, in fact, be the last common ancestor of extant African apes (Kunimatsu Et al. 2007; Mocke Et al. 2002). The previously underappreciated diversity of Late Miocene apes in Africa might be due to poor sampling of the fossil record in Africa.<\/p>\r\n\r\n<h4 class=\"import-Normal\"><em>Eurasian Ape Diversity<\/em><\/h4>\r\n<p class=\"import-Normal\">With the establishment of the <em>Gomphotherium<\/em> Landbridge (a result of the closure of the Eastern Mediterranean seaway; R\u00f6gl 1999), the Middle Miocene was an exciting time for hominoid radiations outside of Africa (see Figure 9.23). Eurasian hominoid species exploited their environments in many different ways in the Miocene. Food exploitation ranged from soft-fruit feeding in some taxa to hard-object feeding in others, in part owing to seasonal fluctuations and the necessary adoptions of fallback foods (DeMiguel Et al. 2014). For example, the molars of <em>Oreopithecus bambolii<\/em> (Family Hominidae) have relatively long lower-molar shearing crests, suggesting that this hominoid was very folivorous (Ungar and Kay 1995). Associated with variation in diet, there is great variation in the degree to which cranial features (e.g., zygomatic bone or supraorbital tori) are developed across the many taxa (Cameron 1997); however, Middle Miocene fossils tend to exhibit relatively thick molar enamel and relatively robust jaws (Andrews and Martin 1991).<\/p>\r\n<p class=\"import-Normal\">In Spain, the cranium with upper dentition, part of a mandible, and partial skeleton of <em>Pliobates <\/em>(Family Pliobatidae), a small-bodied ape (4\u20135 kg; 9\u201311 lbs.), was discovered in deposits dating to 11.6 million years ago (Alba Et al. 2015). It is believed to be a frugivore with a relative brain size that overlaps with modern cercopithecoids. The fossilized postcrania of <em>Pliobates<\/em> suggest that this ape might have had a unique style of locomotion, including the tendency to walk across the branches of trees with its palms facing downward and flexible wrists that permitted rotation of the forearm during climbing. However, the anatomy of the distal humerus differs from those of living apes in ways that suggest that <em>Pliobates<\/em> was less efficient at stabilizing its elbow while suspended (Benefit and McCrossin 2015). Two other recently described apes from Spain, <em>Pierolapithecus <\/em>and <em>Anoiapithecus<\/em>, are known from relatively complete skeletons. <em>Pierolapithecus<\/em> had a very projecting face and thick molar enamel as well as some skeletal features that suggest (albeit controversially) a less suspensory locomotor style than in extant apes (Moy\u00e0-Sol\u00e0 Et al. 2004). In contrast to <em>Pierolapithecus<\/em>, the slightly younger <em>Anoiapithecus<\/em> has a very flat face (Moy\u00e0-Sol\u00e0 Et al. 2009).<\/p>\r\n<p class=\"import-Normal\">Post-cranial evidence for suspensory or well-developed orthograde behaviours in apes does not appear until the Late Miocene of Europe. Primary evidence supporting these specialized locomotor modes includes the relatively short lumbar vertebrae of <em>Oreopithecus <\/em>(Figure 9.26) and <em>Dryopithecus<\/em> (Maclatchy 2004). Further, fossil material of the lower torso of <em>O. bambolii <\/em>(which dates to the <em>Pan<\/em>-hominin divergence) conveys a higher degree of flexion-extension abilities in the lumbar region (lower back) than what is possible in extant apes. Additionally, the hindlimb of <em>O. bambolii <\/em>is suggested to have supported powerful hip adduction during climbing (Hammond Et al. 2020). The Late Miocene saw the extinction of most of the Eurasian hominoids in an event referred to as the Vallesian Crisis (Agust\u00ed Et al. 2003). Among the latest surviving hominoid taxa in Eurasia were <em>Oreopithecus<\/em> and <em>Gigantopithecus<\/em>, the latter of which held out until the Pleistocene in Asia and was probably even sympatric with <em>Homo erectus<\/em> (Cachel 2015).<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"436\"]<img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image8-2-1.jpg\" alt=\"Posterior view of ancient ape skeleton.\" width=\"436\" height=\"775\" \/> Figure 9.26: Skeleton of Oreopithecus bambolii. Credit: <a class=\"rId107\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Oreopithecus_bambolii_1.JPG\">Oreopithecus bambolii 1<\/a> by <a class=\"rId108\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Ghedoghedo\">Ghedoghedo<\/a> is under a <a class=\"rId109\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.[\/caption]\r\n<h3 class=\"import-Normal\"><strong>The Origins of Extant Apes<\/strong><\/h3>\r\n<p class=\"import-Normal\">The fossil record of the extant apes is somewhat underwhelming: it ranges from being practically nonexistent for some taxa (e.g., chimpanzees) to being a little better for others (e.g., humans). There are many possible reasons for these differences in fossil abundance, and many are associated with the environmental conditions necessary for the fossilization of bones. One way to understand the evolution of extant apes that is not so dependent on the fossil record is via molecular evolutionary analyses. This can include counting up the differences in the genetic sequence between two closely related species to estimate the amount of time since these species shared a common ancestor. This is called a molecular clock, and it is often calibrated using fossils of known absolute age that stand in for the last common ancestor of a particular clade. Molecular clock estimates have placed the Hylobatidae and Hominidae split between 19.7 million and 24.1 million years ago, the African ape and Asian ape split between 15.7 million and 19.3 million years ago, and the split of Hylobatidae into its current genera between 6.4 million and 8 million years ago (Israfil Et al. 2011).<\/p>\r\n\r\n<h4 class=\"import-Normal\"><em>Small Ape Origins and Fossils<\/em><\/h4>\r\n<p class=\"import-Normal\">Unfortunately, the fossil record for the small (formerly \u201clesser\u201d) apes is meager, particularly in Miocene deposits. One possible early hylobatid is <em>Laccopithecus robustus<\/em>, a Late Miocene catarrhine from China (Harrison 2016). Although it does share some characteristics with modern gibbons and siamangs (including an overall small body size and a short face), <em>Laccopithecus<\/em> most likely represents a plesiomorphic stem catarrhine and is therefore distantly related to extant apes (Jablonski &amp; Chaplin 2009). A more likely candidate for the hylobatid stem is another Late Miocene taxon from China, <em>Yuanmoupithecus xiaoyuan<\/em>. Interpretation of its phylogenetic standing, however, is complicated by contradicting dental features\u2014some of them quite plesiomorphic\u2014which some believe best place <em>Yuanmoupithecus<\/em> as a stem hylobatid (Harrison 2016). Recently, a Middle Miocene Indian fossil ape, <em>Kapi ramnagarensis<\/em>, has extended the fossil record of small apes by approximately five million years. Its teeth are suggestive of a shift to a more frugivorous diet and it is likely a stem hylobatid (Gilbert Et al. 2020). The history of Hylobatidae becomes clearer in the Pleistocene, with fossils representing extant genera.<\/p>\r\n\r\n<h4 class=\"import-Normal\"><em>Great Ape Origins and Fossils<\/em><\/h4>\r\n<p class=\"import-Normal\">The most extensive fossil record of a modern great ape is that of our own genus, <em>Homo<\/em>. However, the evolutionary history of the Asian great ape, the orangutan (<em>Pongo<\/em>), is becoming clearer. Today, orangutans are found only on the islands of Borneo and Sumatra. However, Pleistocene-aged teeth, attributed to <em>Pongo<\/em>, have been found in Cambodia, China, Laos, Peninsular Malaysia, and Vietnam\u2014demonstrating the vastness of the orangutan\u2019s previous range (Ibrahim Et al. 2013; Wang Et al. 2014). <em>Sivapithecus <\/em>from the Miocene of India and Pakistan is represented by many specimens, including parts of the face. <em>Sivapithecus<\/em> is very similar to <em>Pongo<\/em>, especially in the face, and it probably is closely related to ancestral orangutans (Pilbeam 1982). Originally, jaws and teeth belonging to the former genus <em>Ramapithecus<\/em> were thought to be important in the origin of humans (Simons 1961), but now these are recognized as specimens of <em>Sivapithecus<\/em> (Kelley 2002). Postcranial bones of <em>Sivapithecus<\/em>, however, suggest a more generalized locomotor mode\u2014including terrestrial locomotion\u2014than seen in <em>Pongo <\/em>(Pilbeam Et al. 1990). Stable carbon and oxygen isotope data from dental enamel have reconstructed the paleoecological space of <em>Sivapithecus <\/em>(as well as the contemporaneous Late Miocene pongine <em>Khoratpithecus<\/em>) within the canopies of forested habitats (Habinger Et al. 2022).<\/p>\r\n<p class=\"import-Normal\">A probable close relative of <em>Sivapithecus <\/em>is the amazing <em>Gigantopithecus<\/em> (see Figure 9.24). Known only from teeth and jaws from China and India (e.g., Figure 9.27), this ape probably weighed as much as 270 kg (595 lbs.) and was likely the largest primate ever (Bocherens Et al. 2017). Because of unique features of its teeth (including molarized premolars and patterns of wear) and its massive size, it has been reconstructed as a bamboo specialist, somewhat like the modern panda. Small silica particles (phytoliths) from grasses have been found stuck to the molars of <em>Gigantopithecus<\/em> (Ciochon Et al. 1990). Recent studies evaluating the carbon isotope composition of the enamel sampled from <em>Gigantopithecus<\/em> teeth suggest that this ape exploited a wide range of vegetation, including fruits, leaves, roots, and bamboo (Bocherens Et al. 2017). Its face is reminiscent of that of modern orangutans and it might belong in the same family, Pongidae (Kelley 2002).<\/p>\r\n\r\n\r\n[caption id=\"\" align=\"aligncenter\" width=\"488\"]<img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image12.jpg\" alt=\"Superior view of mandible and teeth.\" width=\"488\" height=\"533\" \/> Figure 9.27: Cast of the mandible of Gigantopithecus blacki. Credit: <a class=\"rId111\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Gigantopithecus%20blacki%20mandible%20010112.jpg\">Gigantopithecus blacki mandible 010112<\/a> by <a class=\"rId112\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Wilson44691\">Wilson44691<\/a> is under a <a class=\"rId113\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.[\/caption]\r\n<p class=\"import-Normal\">In Africa, the first fossil to be confidently attributed to <em>Pan<\/em>, and known to be the earliest evidence of a chimpanzee, was described based on teeth found in Middle Pleistocene deposits in the Eastern Rift Valley of Kenya (McBrearty and Jablonski 2005). Paleoenvironmental reconstructions of this locality suggest that this early chimpanzee was living in close proximity to early <em>Homo<\/em> in a closed-canopy wooded habitat. Similarly, fossil teeth and mandibular remains attributed to two species of Middle-Late Miocene apes\u2014<em>Chororapithecus abyssinicus<\/em> (from Ethiopia; Suwa Et al. 2007) and <em>Nakalipithecus nakayamai<\/em> (from Kenya; Kunimatsu Et al. 2007)\u2014have been suggested as basal members of the gorilla clade.<\/p>\r\n<p class=\"import-Normal\">While the deposits of Eastern Africa have yielded a profound record of our fossil hominin ancestors, the continent\u2019s rainforests remain a \u201cpalaeontological desert\u201d (Rosas Et al. 2022). Clearly, more work is needed to fill in the large gaps in the fossil record of the nonhuman great apes. The twentieth century witnessed the discovery of many hominin fossils in East Africa, which have been critical for improving our understanding of human evolution. While twenty-first-century conservationists fight to prevent the extinction of the living great apes, perhaps efforts by twenty-first-century paleoanthropologists will yield the evolutionary story of these, our closest relatives.<\/p>\r\n\r\n<h2>Cultural Connections and Contemporary Implications<\/h2>\r\nEven today, continued study of primate evolution and the gathering of new evidence are critical to our understanding of our biological history. The Piltdown Man hoax is a striking example of how, without modern carbon dating technology and continued study in the field, false conclusions and evidence can be accepted as truth. In 1912, Charles Dawson\u2014an amateur antiquarian and archeologist\u2014presented a combination of human skull fragments and an ape-like lower jaw as evidence of a previously unknown \u2018missing link\u2019 in the line of human ancestors (Groote et al., 2016). Immediately, members of the British archaeological establishment accepted the specimen because it seemed consistent with the belief that a larger\u2014human-like\u2014brain was among the earliest features to evolve in humans. This specimen remained largely accepted for decades until, in 1953, modern chemical analyses proved that the remains had been deliberately altered: the jaw had been stained, the teeth filed, and the skull fragments were subsequently carbon-dated to only approximately 600 years old (Australian Museum, n.d.).\r\n\r\nAmong many such cases, the Piltdown Man hoax demonstrates why the continued study of primate evolution remains imperative to contemporary biological anthropology. As new discoveries, technologies, and analytical methods provide additional evidence, our understanding of human biological history can be continually evaluated and refined.\r\n\r\nDiscussion questions\r\n<ul>\r\n \t<li style=\"font-weight: 400\">Can you think of an example where a belief that was widely accepted as scientific truth became harmful, not only to science but also to the people within that society?<\/li>\r\n \t<li style=\"font-weight: 400\">Why is it important for scientists to remain open to revising their conclusions when new evidence emerges? How does the Piltdown Man hoax demonstrate the importance of this?<\/li>\r\n<\/ul>\r\n<div class=\"textbox shaded\">\r\n<h2>Summary<\/h2>\r\nWhile there are large gaps in the fossil records linking primates to early hominins, evolutionary trends make it clear that humans are one branch of the broader primate family tree. In this chapter we go over the major development that characterize primate evolution: enhanced vision, grasping hands and feet, greater reliance on social behaviour, and increased brain complexity. It is these traits which distinguish primates from other mammals and furthermore, help define major subsections within the primate class, such as strepsirrhines, haplorhines, monkeys, apes, and ultimately hominins.\r\n\r\nWithin this chapter, we also examine how anthropologists reconstruct these evolutionary relationships. Fossil evidence has provided key information about when and where different primates lived, while genetic data such as skeletal features allow researchers to understand how extinct species moved, ate, and interacted with their environments. Just as crucial are the influences of Earth\u2019s changing environments: continental drift, glacial cycles, and long-term climate shifts have repeatedly reshaped habitats, driving both extinctions and the emergence of new adaptive forms, including the emergence of our own human lineage.\r\n<h2 class=\"import-Normal\">Review Questions<strong>\r\n<\/strong><\/h2>\r\n<ul>\r\n \t<li>Compare three major hypotheses about primate origins, making reference to each one\u2019s key ecological reason for primate uniqueness.<\/li>\r\n \t<li>Explain how changes in temperature, rainfall, and vegetation led to major changes in primate biogeography over the Early Tertiary.<\/li>\r\n \t<li>List some euprimate features that plesiadapiforms have and some that they lack.<\/li>\r\n \t<li>Contrast adapoids and omomyoids in terms of life habits.<\/li>\r\n \t<li>Describe one piece of evidence for each of the adapoid, omomyoid, and tarsier origin hypotheses for anthropoids.<\/li>\r\n \t<li>Discuss the biogeography of the origins of African great apes and orangutans using examples from the Miocene ape fossil record.<\/li>\r\n<\/ul>\r\n<\/div>\r\n<h2 class=\"import-Normal\">Key Terms<strong>\r\n<\/strong><\/h2>\r\n<p class=\"import-Normal\"><strong>Adapoidea<\/strong>: Order: Primates. One of the earliest groups of euprimates (true primates; earliest records from the early Eocene).<\/p>\r\n<p class=\"import-Normal\"><strong>A<\/strong><strong>daptive radiations<\/strong>: Rapid diversifications of single lineages into many species which may present unique morphological features in response to different ecological settings.<\/p>\r\n<p class=\"import-Normal\"><strong>Ancestral traits<\/strong>: Features that were inherited from a common ancestor and which remain (largely) unchanged.<\/p>\r\n<p class=\"import-Normal\"><strong>Anthropoids<\/strong>:Group containing monkeys and apes, including humans.<\/p>\r\n<p class=\"import-Normal\"><strong>Auditory bulla<\/strong>: The rounded bony floor of the middle ear cavity.<\/p>\r\n<p class=\"import-Normal\"><strong>Bilophodonty<\/strong>: Dental condition in which the cusps of molar teeth form ridges (or lophs) separated from each other by valleys (seen, e.g., in modern catarrhine monkeys).<\/p>\r\n<p class=\"import-Normal\"><strong>Catarrhines<\/strong>: Order: Primates; Suborder: Anthropoidea; Infraorder: Catarrhini. Group, with origins in Africa and Asia, that contains monkeys and apes, including humans.<\/p>\r\n<p class=\"import-Normal\"><strong>Clade<\/strong>:Group containing all of the descendants of a single ancestor. A portion of a phylogenetic tree represented as a bifurcation (node) in a lineage and all of the branches leading forward in time from that bifurcation.<\/p>\r\n<p class=\"import-Normal\"><strong>Convergent evolution<\/strong>: The independent evolution of a morphological feature in animals not closely related (e.g., wings in birds and bats).<\/p>\r\n<p class=\"import-Normal\"><strong>Crown<\/strong>: Smallest monophyletic group (clade) containing a specified set of extant taxa and all descendants of their last common ancestor.<\/p>\r\n<p class=\"import-Normal\"><strong>Diastema<\/strong>: Space between adjacent teeth.<\/p>\r\n<p class=\"import-Normal\"><strong>Diffuse coevolution<\/strong>: The ecological interaction between whole groups of species (e.g., primates) with whole groups of other species (e.g., fruiting trees).<\/p>\r\n<p class=\"import-Normal\"><strong>Ectotympanic<\/strong>: Bony ring or tube that holds the tympanic membrane (eardrum).<\/p>\r\n<p class=\"import-Normal\"><strong>Euprimates<\/strong>: Order: Primates. True primates or primates of modern aspect.<\/p>\r\n<p class=\"import-Normal\"><strong>Haplorhines<\/strong>: Group containing catarrhines, platyrrhines, and tarsiers.<\/p>\r\n<p class=\"import-Normal\"><strong>Hominins<\/strong>: Modern humans and any extinct relatives more closely related to us than to chimpanzees.<\/p>\r\n<p class=\"import-Normal\"><strong>Mandibular symphysis<\/strong>: Fibrocartilaginous joint between the left and right mandibular segments, located in the midline of the body.<\/p>\r\n<p class=\"import-Normal\"><strong>Omomyoidea<\/strong>: Order: Primates; Superfamily: Omomyoidea. One of the earliest groups of euprimates (true primates; earliest record in the early Eocene).<\/p>\r\n<p class=\"import-Normal\"><strong>Petrosal bone<\/strong>: The portion of the temporal bone that houses the inner ear apparatus.<\/p>\r\n<p class=\"import-Normal\"><strong>Plagiaulacoid<\/strong>: Dental condition where at least one of the lower cheek-teeth (molars or premolars) is a laterally compressed blade.<\/p>\r\n<p class=\"import-Normal\"><strong>Platyrrhines<\/strong>: Order: Primates; Suborder: Anthropoidea; Infraorder: Platyrrhini. Group containing monkeys found in the Americas.<\/p>\r\n<p class=\"import-Normal\"><strong>Plesiadapiforms<\/strong>: Order: Plesiadapiformes. Archaic primates or primate-like placental mammals (Early Paleocene\u2013Late Eocene).<\/p>\r\n<p class=\"import-Normal\"><strong>P<\/strong><strong>lesiomorphic<\/strong>: Having features that are shared by different groups which arose from a common ancestor.<\/p>\r\n<p class=\"import-Normal\"><strong>Stem<\/strong>: Taxa that are basal to a given crown group but are more closely related to the crown group than to the closest living sister taxon of the crown group.<\/p>\r\n<p class=\"import-Normal\"><strong>Strepsirrhines<\/strong>: Order: Primates; Suborder: Stresirrhini. Group containing lemurs, lorises, and galagos (does not include tarsiers).<\/p>\r\n<p class=\"import-Normal\"><strong>Toothcomb<\/strong>: Dental condition found in modern strepsirrhines in which the lower incisors and canines are laterally compressed and protrude forward at a nearly horizontal inclination. This structure is used in grooming.<\/p>\r\n\r\n<\/div>\r\n<div class=\"__UNKNOWN__\">\r\n<h2 class=\"import-Normal\">For Further Exploration<strong>\r\n<\/strong><\/h2>\r\n<p class=\"import-Normal\">Beard, Chris. 2004. <em>The Hunt for the Dawn Monkey: Unearthing the Origins of Monkeys, Apes, and Humans<\/em>. Berkeley: University of California Press.<\/p>\r\n<p class=\"import-Normal\">Begun, David R. 2010. \u201cMiocene Hominids and the Origins of the African Apes and Humans.\u201d <em>Annual Review of Anthropology<\/em> 39: 67\u201384.<\/p>\r\n<p class=\"import-Normal\">Fleagle, John G. 2013. <em>Primate Adaptation and Evolution.<\/em> Third edition. San Diego, CA: Academic Press.<\/p>\r\n<p class=\"import-Normal\">Gebo, Daniel L., ed. 1993. <em>Postcranial Adaptations in Nonhuman Primates<\/em>. Dekalb: Northern Illinois University Press.<\/p>\r\n<p class=\"import-Normal\">Godfrey, Laurie R., and William L. Jungers. 2002. \u201cQuaternary Fossil Lemurs.\u201d In <em>The Primate Fossil Record, <\/em>edited by Walter C. Hartwig, 97\u2013121. Cambridge: Cambridge University Press.<\/p>\r\n<p class=\"import-Normal\">Godinot, Marc. 2006. \u201cLemuriform Origins as Viewed from the Fossil Record.\u201d <em>Folia Primatologica<\/em> 77 (6): 446\u2013464.<\/p>\r\n<p class=\"import-Normal\">Kay, Richard F. 2018. \u201c100 Years of Primate Paleontology.\u201d <em>American Journal of Physical Anthropology<\/em> 165 (4): 652\u2013676.<\/p>\r\n<p class=\"import-Normal\">Marivaux, Laurent. 2006. \u201cThe Eosimiid and Amphipithecid Primates (Anthropoidea) from the Oligocene of the Bugti Hills (Balochistan, Pakistan): New Insight into Early Higher Primate Evolution in South Asia.\u201d <em>Palaeovertebrata, Montpellier <\/em>34 (1\u20132): 29\u2013109.<\/p>\r\n<p class=\"import-Normal\">Martin, R. D. 1990. <em>Primate Origins and Evolution<\/em><em>: A <\/em><em>Phylogenetic Reconstruction<\/em>. Princeton: Princeton University Press.<\/p>\r\n<p class=\"import-Normal\">Rose, Kenneth D., Marc Godinot, and Thomas M. Bown. 1994. \u201cThe Early Radiation of Euprimates and the Initial Diversification of Omomyidae.\u201d In <em>Anthropoid Origins: The Fossil Evidence, <\/em>edited by John G. Fleagle and Richard F. Kay, 1\u201328. New York: Plenum Press.<\/p>\r\n<p class=\"import-Normal\">Ross, Callum F. 1999. \u201cHow to Carry Out Functional Morphology.\u201d <em>Evolutionary Anthropology<\/em> 7 (6): 217\u2013222.<\/p>\r\n<p class=\"import-Normal\">Seiffert, Erik R. 2012. \u201cEarly Primate Evolution in Afro-Arabia.\u201d Evolutionary Anthropology: Issues, News, and Reviews 21(6): 239\u2013253.<\/p>\r\n<p class=\"import-Normal\">Szalay, Frederic S., and Eric Delson. 1979. Evolutionary History of the Primates. New York: Academic Press.<\/p>\r\n<p class=\"import-Normal\">Ungar, Peter S. 2002. \u201cReconstructing the Diets of Fossil Primates.\u201d In <em>Reconstructing Behavior in the Primate Fossil Record<\/em>, edited by Joseph Plavcan, Richard F. Kay, William Jungers, and Carel P. van Schaik, 261\u2013296. New York: Kluwer Academic\/Plenum Publishers.<\/p>\r\n\r\n<h2 class=\"import-Normal\">References<\/h2>\r\n<p class=\"import-Normal\">Agust\u00ed, J., A. Sanz de Siria, and M. Garc\u00e9s M. 2003. \u201cExplaining the End of the Hominoid Experiment in Europe.\u201d <em>Journal of Human Evolution<\/em> 45 (2): 145\u2013153.<\/p>\r\n<p class=\"import-Normal\">Alba, David M., Sergio Alm\u00e9cija, Daniel DeMiguel, Josep Fortuny, Miriam P\u00e9rez de los R\u00edos, Marta Pina, Josep M. 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Rensberger. 1983. \u201cUpper Dentition of <em>Ekgmowechashala<\/em> (Omomyoid Primate) from the John Day Formation, Oligo-Miocene of Oregon.\u201d <em>Folia Primatologica<\/em> 41(1-2): 102\u2013111.<\/p>\r\n<p class=\"import-Normal\">Rosenberger, Alfred L. 2010. \u201cPlatyrrhines, PAUP, Parallelism, and the Long Lineage Hypothesis: A Reply to Kay <em>et al. <\/em>(2008).\u201d <em>Journal of Human Evolution<\/em> 59 (2): 214\u2013217.<\/p>\r\n<p class=\"import-Normal\">Ross, Callum F. 2000. \u201cInto the Light: The Origins of Anthropoidea.\u201d <em>Annual Review of Anthropology<\/em> 29: 147\u2013194.<\/p>\r\n<p class=\"import-Normal\">Ross, Callum F., and Richard F. Kay, eds. 2004. <em>Anthropoid Origins: New Visions<\/em>. 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Simons. 1996. \u201cDental Microwear and Microstructure in Early Oligocene Primates from the Fayum, Egypt: Implications for Diet.\u201d <em>American Journal of Physical Anthropology<\/em> 101 (4): 527\u2013543.<\/p>\r\n<p class=\"import-Normal\">Ungar, Peter S., and Richard F. Kay. 1995. \u201cThe Dietary Adaptations of European Miocene Catarrhines.\u201d <em>Proceedings of the National Academy of Sciences of the United States of America<\/em> 92 (12): 5479\u20135481.<\/p>\r\nUngar, Peter S. 2002. \u201cReconstructing the Diets of Fossil Primates.\u201d In Reconstructing Behaviour in the Primate Fossil Record, edited by Joseph Plavcan, Richard F. Kay, William Jungers, and Carel P. van Schaik, 261\u2013296. New York: Kluwer Academic\/Plenum Publishers.\r\n<p class=\"import-Normal\">Wang, Cui-Bin, Ling-Xia Zhao, Chang-Zhu Jin, Yuan Wang, Da-Gong Qin, and Wen-Shi Pan. 2014. \u201cNew Discovery of Early Pleistocene Orangutan Fossils from Sanhe Cave in Chongzuo, Guangxi, Southern China.\u201d <em>Quaternary International<\/em> 354: 68\u201374.<\/p>\r\n<p class=\"import-Normal\">Ward, C. V., A. Walker, and M. F. Teaford. 1991. \u201c<em>Proconsul<\/em> Did Not Have a Tail.\u201d <em>Journal of Human Evolution<\/em> 21 (3): 215\u2013220.<\/p>\r\n<p class=\"import-Normal\">Wheeler, Brandon C. 2010. \u201cCommunity Ecology of the Middle Miocene Primates of La Venta, Colombia: The Relationship between Ecological Diversity, Divergence Time, and Phylogenetic Richness.\u201d <em>Primates<\/em> 51 (2): 131\u2013138.<\/p>\r\n<p class=\"import-Normal\">Williams, Blythe A., and Richard F. Kay. 1995. \u201cThe Taxon Anthropoidea and the Crown Clade Concept.\u201d <em>Evolutionary Anthropology<\/em> 3 (6): 188\u2013190.<\/p>\r\n<p class=\"import-Normal\">Williams, Blythe A., Richard F. Kay, and E. Christopher Kirk. 2010a. \u201cNew Perspectives on Anthropoid Origins.\u201d <em>Proceedings of the National Academy<\/em> <em>of the United States of America<\/em> 107 (11): 4797\u20134804.<\/p>\r\n<p class=\"import-Normal\">Williams, Blythe A., Richard F. Kay, E. Christopher Kirk, and Callum F. Ross. 2010b. \u201c<em>Darwinius masillae<\/em> Is a European Middle Eocene Stem Strepsirrhine\u2014A Reply to Franzen et al.\u201d <em>Journal of Human Evolution<\/em> 59(5): 567\u2013573.<\/p>\r\n<p class=\"import-Normal\">Wilson Mantilla, G. P., S. G. B. Chester, W. A. Clemens, J. R. Moore, C. J. Sprain, B. T. Hovatter, W. S. Mitchell, W. W. Mans, R. Mundil, and P. R. Renne. 2021. \u201cEarliest Palaeocene Purgatoriids and the Initial Radiation of Stem Primates.\u201d <em>Royal Society Open Science<\/em> 8(2):210050. doi:10.1098\/rsos.210050.<\/p>\r\n\r\n<h2 class=\"import-Normal\">Acknowledgements<\/h2>\r\n<p class=\"import-Normal\">We are immensely grateful to the editors of this book, Drs. Beth Shook, Lara Braff, Katie Nelson, and Kelsie Aguilera, for their time and commitment to making this knowledge freely accessible to all, and for giving us the opportunity to participate in this important project.<\/p>\r\n\r\n<\/div>\r\n<\/div>","rendered":"<div class=\"__UNKNOWN__\">\n<div class=\"textbox\">\n<p>Learn more about <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/about-the-authors\/\">authors and editors<\/a>,<a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/student-contributors\/\"> Hess&#8217; student contributions<\/a>, and <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/versioning-history\/\">versioning history<\/a>.<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\">Learning Objectives<\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li>Understand the major trends in primate evolution from the origin of primates to the origin of our own species.<\/li>\n<li>Learn about primate adaptations and how they characterize major primate groups.<\/li>\n<li>Discuss the kinds of evidence that anthropologists use to find out how extinct primates are related to each other and to living primates.<\/li>\n<li>Recognize how the changing geography and climate of Earth have influenced where and when primates have thrived or gone extinct.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p class=\"import-Normal\">The first fifty million years of primate evolution was a series of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1683\">adaptive radiations<\/a><\/strong> leading to the diversification of the earliest lemurs, monkeys, and apes. The primate story begins in the canopy and understory of conifer-dominated forests, with our small, furtive ancestors subsisting at night, beneath the notice of day-active dinosaurs.<\/p>\n<p class=\"import-Normal\">From the ancient <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1684\"><strong>plesiadapiforms<\/strong><\/a> (archaic primates) to the earliest groups of true primates (<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1686\"><strong>euprimates<\/strong><\/a>) (Bloch and Boyer 2002), the origin of our own order is characterized by the struggle for new food sources and microhabitats in the arboreal setting. Climate change forced major extinctions as the northern continents became increasingly dry, cold, and seasonal and as tropical rainforests gave way to deciduous forests, woodlands, and eventually grasslands. Lemurs, lorises, and tarsiers\u2014once diverse groups containing many species\u2014became rare, except for lemurs in Madagascar, where there were no anthropoid competitors and perhaps few predators. Meanwhile, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1685\">anthropoids<\/a><\/strong> (monkeys and apes) likely emerged in Asia and then dispersed across parts of the northern hemisphere, Africa, and ultimately South America. The movement of continents, shifting sea levels, and changing patterns of rainfall and vegetation contributed to the developing landscape of primate biogeography, morphology, and behaviour. Today\u2019s primates provide modest reminders of the past diversity and remarkable adaptations of their extinct relatives. This chapter explores the major trends in primate evolution from the origin of the Order Primates to the beginnings of our own lineage, providing a window into these stories from our ancient past.<\/p>\n<h2 class=\"import-Normal\">Major Hypotheses About Primate Origins<\/h2>\n<p class=\"import-Normal\">For many groups of mammals, there is a key feature that led to their success. A good example is powered flight in bats. Primates lack a feature like this (see Chapter 6). Instead, if there is something unique about primates, it is probably a group of features rather than one single thing. Because of this, anthropologists and palaeontologists struggle to describe an ecological scenario that could explain the rise and success of our own order. Three major hypotheses have been advanced to consider the origin of primates and to explain what makes our order distinct among mammals (Figure 9.1); these are described below.<\/p>\n<figure id=\"attachment_277\" aria-describedby=\"caption-attachment-277\" style=\"width: 634px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" class=\"wp-image-255\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2023\/03\/8.1.jpg\" alt=\"Primates swinging in tree, eating an insect, and eating fruit.\" width=\"634\" height=\"221\" \/><figcaption id=\"caption-attachment-277\" class=\"wp-caption-text\">Figure 9.1: The three major hypotheses are (a) the arboreal hypothesis, (b) the visual predation hypothesis, and (c) the angiosperm-primate coevolution hypothesis. Credit: Primate origin hypotheses original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by <a class=\"rId13\" href=\"https:\/\/marynelsonstudio.com\">Mary Nelson<\/a> is under a <a class=\"rId14\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Arboreal Hypothesis<\/strong><\/h3>\n<p class=\"import-Normal\">In the 1800s, many anthropologists viewed all animals in relation to humans. That is, animals that were more like humans were considered to be more \u201cadvanced\u201d and those lacking humanlike features were considered more \u201cprimitive.\u201d This way of thinking was particularly obvious in studies of primates. A more modern way of referring to members of a group that lack certain evolutionary innovations seen in other members is to call them <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1688\"><strong>plesiomorphic<\/strong><\/a> (literally \u201canciently shaped\u201d). The state of their morphological features is sometimes referred to as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1689\"><strong>ancestral<\/strong><strong> traits<\/strong><\/a>.<\/p>\n<p class=\"import-Normal\">Thus, when anthropologists sought features that separate primates from other mammals, they focused on features that were least developed in lemurs and lorises, more developed in monkeys, and most developed in apes (Figure 9.2). Frederic Wood Jones, one of the leading anatomist-anthropologists of the early 1900s, is usually credited with the Arboreal Hypothesis of primate origins (Jones 1916). This hypothesis holds that many of the features of primates evolved to improve locomotion in the trees; this way of getting around is referred to as arboreal. For example, the grasping hands and feet of primates are well suited to gripping tree branches of various sizes and our flexible joints are good for reorienting the extremities in many different ways. A mentor of Jones, Grafton Elliot Smith, had suggested that the reduced olfactory system, acute vision, and forward-facing eyes of primates are adaptations for making accurate leaps and bounds through a complex, three-dimensional canopy (Smith 1912). The forward orientation of the eyes in primates causes the visual fields to overlap, enhancing depth perception, especially at close range. Evidence to support this hypothesis includes the facts that many extant primates are arboreal, and the plesiomorphic members of most primate groups are dedicated arborealists. The Arboreal Hypothesis was well accepted by most anthropologists at the time and for decades afterward.<\/p>\n<figure style=\"width: 663px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image27-2.png\" alt=\"Diagram shows primates descended from Plesiadapiforms.\" width=\"663\" height=\"543\" \/><figcaption class=\"wp-caption-text\">Figure 9.2: Primate family tree showing major groups. Disconnected lines show uncertainty about relationships. Two lines lead to tarsiers from different possible groups of origin. <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 class=\"rId16\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Primate family tree (Figure 8.2)<\/a> by Jonathan M. G. Perry 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<h3 class=\"import-Normal\"><strong>Visual Predation Hypothesis<\/strong><\/h3>\n<p class=\"import-Normal\">In the late 1960s and early 1970s, Matt Cartmill studied and tested the idea that the characteristic features of primates evolved in the context of arboreal locomotion. Cartmill noted that squirrels climb trees (and even vertical walls) very effectively, even though they lack some of the key adaptations of primates. As members of the Order Rodentia, squirrels also lack the hand and foot anatomy of primates. They have claws instead of flattened nails and their eyes face more laterally than those of primates. Cartmill reasoned that there must be some other explanation for the unique traits of primates. He noted that some nonarboreal animals share at least some of these traits with primates; for example, cats and predatory birds have forward-facing eyes that enable visual field overlap. Cartmill suggested that the unique suite of features in primates is an adaptation to detecting insect prey and guiding the hands (or feet) to catch insects (Cartmill 1972). His hypothesis emphasizes the primary role of vision in prey detection and capture; it is explicitly comparative, relying on form-function relationships in other mammals and nonmammalian vertebrates. According to Cartmill, many of the key features of primates evolved for preying on insects in this special manner (Cartmill 1974).<\/p>\n<h3 class=\"import-Normal\"><strong>Angiosperm-Primate Coevolution Hypothesis<\/strong><\/h3>\n<p class=\"import-Normal\">The visual predation hypothesis was unpopular with some anthropologists. One reason for this is that many primates today are not especially predatory. Another is that, whereas primates do seem well adapted to moving around in the smallest, terminal branches of trees, insects are not necessarily easier to find there. A counterargument to the visual predation hypothesis is the angiosperm-primate coevolution hypothesis. Primate ecologist Robert Sussman (Sussman 1991) argued that the few primates that eat mostly insects often catch their prey on the ground rather than in tree branches. Furthermore, predatory primates often use their ears more than their eyes to detect prey. Finally, most early primate fossils show signs of having been omnivorous rather than insectivorous. Instead, he argued, the earliest primates were probably seeking fruit. Fruit (and flowers) of angiosperms (flowering plants) often develop in the terminal branches. Therefore, any mammal trying to access those fruits must possess anatomical traits that allow them to maintain their hold on thin branches and avoid falling while reaching for the fruits. Primates likely evolved their distinctive visual traits and extremities in the Paleocene (approximately 65 million to 54 million years ago) and Eocene (approximately 54 million to 34 million years ago) epochs, just when angiosperms were going through a revolution of their own\u2014the evolution of large, fleshy fruit that would have been attractive to a small arboreal mammal. Sussman argued that, just as primates were evolving anatomical traits that made them more efficient fruit foragers, angiosperms were also evolving fruit that would be more attractive to primates to promote better seed dispersal. This mutually beneficial relationship between the angiosperms and the primates was termed coevolution or more specifically <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1691\"><strong>diffuse coevolution<\/strong>.<\/a><\/p>\n<p class=\"import-Normal\">At about the same time, D. Tab Rasmussen noted several parallel traits in primates and the South American woolly opossum, <em>Caluromys<\/em>. He argued that early primates were probably foraging on both fruits and insects (Rasmussen 1990). As is true of <em>Caluromys<\/em> today, early primates probably foraged for fruits in the terminal branches of angiosperms, and they probably used their visual sense to aid in catching insects. Insects are also attracted to fruit (and flowers), so these insects represent a convenient opportunity for a primarily fruit-eating primate to gather protein. This solution is a compromise between the visual predation hypothesis and the angiosperm-primate coevolution hypothesis. It is worth noting that other models of primate origins have been proposed, and these include the possibility that no single ecological scenario can account for the origin of primates.<\/p>\n<h2 class=\"import-Normal\">The Origins of Primates<\/h2>\n<h3 class=\"import-Normal\"><strong>Paleocene: Mammals in the Wake of Dinosaur Extinctions<\/strong><\/h3>\n<p class=\"import-Normal\">Placental mammals, including primates, originated in the Mesozoic Era (approximately 251 million to 65.5 million years ago), the Age of Dinosaurs. During this time, most placental mammals were small, probably nocturnal, and probably avoided predators via camouflage and slow, quiet movement. It has been suggested that the success and diversity of the dinosaurs constituted a kind of ecological barrier to Mesozoic mammals. The extinction of the dinosaurs (and many other organisms) at the end of the Cretaceous Period (approximately 145.5\u201365.5 million years ago) might have opened up these ecological niches, leading to the increased diversity and disparity in mammals of the Tertiary Period (approximately 65.5\u20132.5 million years ago).<\/p>\n<p class=\"import-Normal\">The Paleocene was the first epoch in the Age of Mammals. Soon after the Cretaceous-Tertiary (K-T) extinction event, new groups of placental mammals appear in the fossil record. Many of these groups achieved a broad range of sizes and lifestyles as well as a great number of species before declining sometime in the Eocene (or soon thereafter). These groups were ultimately replaced by the modern orders of placental mammals (Figure 9.3). It is unknown whether these replacements occurred gradually, for example by competitive exclusion, or rapidly, perhaps by sudden geographic dispersals with replacement. In some senses, the Paleocene might have been a time of recovery from the extinction event; it was cooler and more seasonal globally than the subsequent Eocene.<\/p>\n<figure style=\"width: 628px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image26.jpg\" alt=\"Person in front of a mural depicting forest animals.\" width=\"628\" height=\"511\" \/><figcaption class=\"wp-caption-text\">Figure 9.3: A mural of Eocene flora and fauna in North America. Credit: <a class=\"rId19\" href=\"https:\/\/flickr.com\/photos\/126377022@N07\/18404106406\">Image from page 27 of &#8220;Annual report for the year ended June 30 &#8230;&#8221; (1951)<\/a> by <a class=\"rId20\" href=\"https:\/\/flickr.com\/photos\/internetarchivebookimages\/\">Internet Archive Book Images<\/a> has been designated to the <a class=\"rId21\" href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/\">public domain (CC0)<\/a>. This photograph of the mural &#8220;Fauna and flora of middle Eocene in the Wyoming area&#8221; by Jay Matternes, was originally published by the <a class=\"rId22\" href=\"https:\/\/www.si.edu\/\">Smithsonian<\/a>, and can be viewed in context in the <a class=\"rId23\" href=\"https:\/\/archive.org\/details\/annualreportfory1961united\/page\/7\/mode\/1up?view=theater\">online version of this book<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Plesiadapiforms, the Archaic Primates<\/strong><\/h3>\n<p class=\"import-Normal\">The Paleocene epoch saw the emergence of several families of mammals that have been implicated in the origin of primates. These are the plesiadapiforms, which are archaic primates, meaning they possessed some primate features and lacked others. The word <em>plesiadapiform <\/em>means \u201calmost adapiform,\u201d a reference to some similarities between some plesiadapiforms and some adapiforms (or adapoids; later-appearing true primates)\u2014mainly in the molar teeth. Because enamel fossilizes better than other parts of the body, the molar teeth are the parts most often found and first discovered for any new species. Thus, dental similarities were often the first to be noticed by early mammalian paleontologists, partly explaining why plesiadapiforms were thought to be primates. Major morphological differences between plesidapiforms and euprimates (true primates) were observed later when more parts of plesiadapiform skeletons were discovered. Many plesiadapiforms have unusual anterior teeth and most have digits possessing claws rather than nails. So far, no plesiadapiform ever discovered has a postorbital bar (seen in extant <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1712\">strepsirrhines<\/a><\/strong>) or septum (as seen in <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1713\">haplorhines<\/a><\/strong>), and whether or not the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1711\">auditory bulla<\/a><\/strong> was formed by the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1714\"><strong>petrosal bone<\/strong> <\/a>remains unclear for many plesiadapiform specimens. Nevertheless, there are compelling reasons (partly from new skeletal material) for including plesiadapiforms within the Order Primates.<\/p>\n<h4 class=\"import-Normal\"><em>Geographic and Temporal Distribution<\/em><\/h4>\n<p class=\"import-Normal\"><em>Purgatorius<\/em> is generally considered to be the earliest primate. This Paleocene mammal is known from teeth that are very plesiomorphic for a primate. It has some characteristics that suggest it is a basal plesiadapiform, but there is very little to link it specifically with euprimates (see Clemens 2004). Its ankle bones suggest a high degree of mobility, signaling an arboreal lifestyle (Chester Et al. 2015). <em>Purgatorius<\/em> is plesiomorphic enough to have given rise to all primates, including the plesiadapiforms. However, new finds suggest that this genus was more diverse and had more differing tooth morphologies than previously appreciated (Wilson Mantilla Et al. 2021). Plesiadapiform families were numerous and diverse during parts of the Paleocene in western North America and western Europe, with some genera (e.g., <em>Plesiadapis<\/em>; see Figure 9.4) living on both continents (Figure 9.5). Thus, there were probably corridors for plesiadapiform dispersal between the two continents, and it stands to reason that these mammals were living all across North America, including in the eastern half of the continent and at high latitudes. A few plesiadapiforms have been described from Asia (e.g., <em>Carpocristes<\/em>), but the affinities of these remain uncertain.<\/p>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 473.25pt\">\n<caption>Figure 9.4: Families of plesiadapiforms with example genera and traits: a table. Credit: Plesiadapiforms table original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Jonathan M. G. Perry and Stephanie L. Canington is under a <a class=\"rId24\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Content derived from Fleagle 2013.<\/caption>\n<thead>\n<tr style=\"height: 25pt\">\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Family<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Genera<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Morphology<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Location<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Age<\/strong><sup><strong>1<\/strong><\/sup><\/p>\n<p>&nbsp;<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table1-R\" style=\"height: 17pt\">\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Paromomyidae<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Ignacius<\/em><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Long, dagger-like, lower incisor.<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">North America and Europe<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Early Paleocene to Late Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 18pt\">\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Carpolestidae<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Carpolestes<\/em><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Plagiaulacoid dentition. Limb adaptations to terminal branch feeding. Grasping big toe.<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">North America, Europe, and Asia<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Middle Paleocene to Early Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 16pt\">\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Plesiadapidae<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Plesiadapis<\/em><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Mitten-like upper incisor. Diastema. Large body size for group.<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">North America and Europe<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Middle Paleocene to Early Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 1pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\" colspan=\"4\">\n<p class=\"import-Normal\"><sup>1<\/sup> Derived from Fleagle 2013.<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\">\n<p class=\"import-Normal\">\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<figure id=\"attachment_277-2\" aria-describedby=\"caption-attachment-277-2\" style=\"width: 555px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" class=\"wp-image-258\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5-5-e1691791897574.png\" alt=\"Global map with not fully formed continents.\" width=\"555\" height=\"308\" \/><figcaption id=\"caption-attachment-277-2\" class=\"wp-caption-text\">Figure 9.5: Map of the world in the Paleocene, highlighting plesiadapiform localities on lands that would become North America, southern Europe, and eastern Asia. Credit: <a class=\"rId26\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Paleocene Map with Plesiadapiform Localities (Figure 8.4)<\/a> original to<a class=\"rId27\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"> Expl<\/a><a class=\"rId28\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">orations: An Open Invitation to Biological Anthropology<\/a> by Elyssa Ebding at <a class=\"rId29\" href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\">GeoPlace, California State University, Chico<\/a> is under a <a class=\"rId30\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Localities based on Fleagle 2013, 211.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>General Morphological Features<\/em><\/h4>\n<p class=\"import-Normal\">Although there is much morphological variation among the families of plesiadapiforms, some common features unite the group. Most plesiadapiforms were small, the largest being about three kilograms (approximately 7 lbs.; <em>Plesiadapis cookei<\/em>). They had small brains and fairly large snouts, with eyes that faced more laterally than in euprimates. Many species show reduction and\/or loss of the canine and anterior premolars, with the resulting formation of a rodent-like <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1172\">diastema<\/a> <\/strong>(a pronounced gap between the premolars and the incisors, with loss of at least the canine); this probably implies a herbivorous diet. Some families appear to have had very specialized diets, as suggested by unusual tooth and jaw shapes.<\/p>\n<p class=\"import-Normal\">Arguably the most interesting and unusual family of plesiadapiforms is the Carpolestidae. They are almost exclusively from North America (with a couple of possible members from Asia), and mainly from the Middle and Late Paleocene. Their molars are not very remarkable, being quite similar to those of some other plesiadapiforms (e.g., Plesiadapidae). However, their lower posterior premolars (p4) are laterally compressed and blade-like with vertical serrations topped by tiny cuspules. This unusual dental morphology is termed <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1693\"><strong><em>plagiaulacoid<\/em><\/strong> <\/a> (Simpson 1933). The upper premolar occlusal surfaces are broad and are covered with many small cuspules; the blade-like lower premolar might have cut across these cuspules, between them, or both.<\/p>\n<div class=\"mceTemp\"><\/div>\n<figure style=\"width: 314px\" class=\"wp-caption alignleft\"><img loading=\"lazy\" decoding=\"async\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image1-5.png\" alt=\"Small brown animal with long tail.\" width=\"314\" height=\"178\" \/><figcaption class=\"wp-caption-text\">Figure 9.6: An artistic rendition of Carpolestes simpsoni moving along a small diameter support. Credit: <a class=\"rId32\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:CarpolestesCL.png\">CarpolestesCL<\/a> by <a class=\"rId33\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Sisyphos23\">Sisyphos23<\/a> is under a <a class=\"rId34\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Many plesiadapiforms have robust limb bones with hallmarks of arboreality. Instead of having nails, most taxa had sharp claws on most or all of the digits. The extremities show grasping abilities comparable to those of primates and some arboreal marsupials. Nearly complete skeletons have yielded a tremendous wealth of information on locomotor and foraging habits. Many plesiadapiforms appear to have been able to cling to vertical substrates (like a broad tree trunk) using their sharp claws, propelling themselves upward using powerful hindlimbs, bounding along horizontal supports, grasping smaller branches, and moving head-first down tree trunks. In carpolestids in particular, the skeleton appears to have been especially well adapted to moving slowly and carefully in small terminal branches (Figure 9.6).<\/p>\n<\/div>\n<div class=\"textbox shaded\">\n<h3 class=\"import-Normal\">Dig Deeper: Debate on the Relationship of Plesiadapiforms to True Primates<\/h3>\n<p class=\"import-Normal\">In the middle of the twentieth century, treeshrews (Order Scandentia) were often considered part of the Order Primates, based on anatomical similarities between some treeshrews and primates. For many people, plesiadapiforms represented intermediates between primates and treeshrews, so plesiadapiforms were included in Primates as well.<\/p>\n<p class=\"import-Normal\">Studies of reproduction and brain anatomy in treeshrews and lemurs suggested that treeshrews are not primates (e.g., Martin 1968). This was soon followed by the suggestion to also expel plesiadapiforms (Martin 1972) from the Order Primates. Like treeshrews, plesiadapiforms lack a postorbital bar, nails, and details of the ear region that characterize true primates. Many paleoanthropologists were reluctant to accept this move to banish plesiadapiforms (e.g., F. S. Szalay, P. D. Gingerich).<\/p>\n<p class=\"import-Normal\">Later, K. Christopher Beard (1990) found that in some ways, the digits of paromomyid plesiadapiforms are actually more similar to those of dermopterans (Order Dermoptera), the closest living relatives of primates, than they are to those of primates themselves (but see Krause 1991). At the same time, Richard Kay and colleagues (1990) found that cranial circulation patterns and auditory bulla morphology in the paromomyid, <em>Ignacius <\/em>(see Figure 9.4), are more like those of dermopterans than of primates.<\/p>\n<p class=\"import-Normal\">For many anthropologists, this one-two punch effectively removed plesiadapiforms from the Order Primates. In the last two decades, the tide of opinion has turned again, with many researchers reinstating plesiadapiforms as members of the Order Primates. New and more complete specimens demonstrate that the postcranial skeletons of plesiadapiforms, including the hands and feet, were primate-like, not dermorpteran-like (Bloch and Boyer 2002, 2007). New fine-grained CT scans of relatively complete plesiadapiform skulls revealed that they share some key traits with primates to the exclusion of other placental mammals (Bloch and Silcox 2006). Most significant was the suggestion that <em>Carpolestes simpsoni <\/em>possessed an auditory bulla formed by the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1696\"><strong>petrosal <\/strong><strong>bone<\/strong><\/a>, like in all living primates.<\/p>\n<p class=\"import-Normal\">The debate about the status of plesiadapiforms continues, owing to a persistent lack of key bones in some species and owing to genuine complexity of the anatomical traits involved. Maybe plesiadapiforms were the ancestral stock from which all primates arose, with some plesiadapiforms (e.g., carpolestids) nearer to the primate <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1723\">stem<\/a><\/strong> than others.<\/p>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<h3 class=\"import-Normal\"><strong>Adapoids and Omomyoids, the First True Primates<\/strong><\/h3>\n<h4 class=\"import-Normal\"><em>Geographic and Temporal Distribution<\/em><\/h4>\n<p class=\"import-Normal\">The first universally accepted fossil primates are the adapoids (Superfamily <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1695\"><strong>Adapoidea<\/strong><\/a>) and the omomyoids (Superfamily <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1694\">Omomyoidea<\/a>)<\/strong>. These groups become quite distinct over evolutionary time, filling mutually exclusive niches for the most part. However, the earliest adapoids are very similar to the earliest omomyoids.<\/p>\n<p class=\"import-Normal\">The adapoids were mainly diurnal and herbivorous, with some achieving larger sizes than any plesiadapiforms (10 kg; 22 lbs.). By contrast, the omomyoids were mainly nocturnal, insectivorous and frugivorous, and small.<\/p>\n<p class=\"import-Normal\">Both groups appear suddenly at the start of the Eocene, where they are present in western North America, western Europe, and India (Figure 9.7). This wide dispersal of early primates was probably due to the presence of rainforest corridors extending far into northern latitudes.<\/p>\n<figure id=\"attachment_277-3\" aria-describedby=\"caption-attachment-277-3\" style=\"width: 539px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" class=\"wp-image-260\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image22-3-e1691792023503.png\" alt=\"Global map with not fully formed continents and omomyoid localities.\" width=\"539\" height=\"317\" \/><figcaption id=\"caption-attachment-277-3\" class=\"wp-caption-text\">Figure 9.7: Map of the world in the Eocene, highlighting adapoid and omomyoid localities on lands that would become North America, southern Europe, Africa, and Asia. Credit: <a class=\"rId36\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Eocene Map with Adapoid and Omomyoid Localities (Figure 8.6)<\/a> original to <a class=\"rId37\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Elyssa Ebding at <a class=\"rId38\" href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\">GeoPlace, California State University, Chico<\/a> is under a <a class=\"rId39\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Localities based on Fleagle 2013, 229.<\/figcaption><\/figure>\n<p class=\"import-Normal\">In North America and Europe, both groups achieved considerable diversity in the Middle Eocene, then mostly died out at the end of that epoch (Figure 9.8). In some Eocene rock formations in the western United States, adapoids and omomyoids make up a major part of the mammalian fauna. The Eocene of India has yielded a modest diversity of euprimates, some of which are so plesiomorphic that it is difficult to know whether they are adapoids or omomyoids (or even early anthropoids).<\/p>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 473.25pt\">\n<caption>Figure 9.8: Families of adapoids and omomyoids with example genera and traits: a table. Credit: Adapoids and omomyoids table original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Jonathan M. G. Perry and Stephanie L. Canington is under a <a class=\"rId40\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Content derived from Fleagle 2013.<\/caption>\n<thead>\n<tr style=\"height: 25pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Family<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Genera<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Morphology<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Location<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Age<\/strong><sup><strong>1<\/strong><\/sup><\/p>\n<p>&nbsp;<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table2-R\" style=\"height: 18pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Cercamoniidae<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Donrussellia<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Variable in tooth number and jaw shape.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Europe and Asia<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Early to Late Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 16pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Asiadapidae<sup>2<\/sup><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Asiadapis<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Plesiomorphic teeth and jaw resemble early Omomyids.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Asia<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Early Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 16pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Caenopithecidae<sup>3<\/sup><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Darwinius<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Robust jaws with crested molars. Fewer premolars.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Europe, Africa, North America, and Asia<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Middle to Late Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 16pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Adapidae<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Adapis<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Fused mandible. Long molar crests. Large size and large chewing muscles.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Europe<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Late Eocene to Early Oligocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 16pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Sivaladapidae<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Sivaladapis<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Some large with robust jaws.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Asia<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Middle Eocene to Late Miocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 16pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Notharctidae<sup>4<\/sup><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Notharctus<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Canine sexual dimorphism. Lemur-like skull. Clinging and leaping adaptations.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">North America and Europe<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Early to Middle Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 16pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Omomyidae<sup>5<\/sup><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Teilhardina<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Small, nocturnal, frugivorous or insectivorous. Tarsier-like skull in some.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">North America, Europe, and Asia<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Early Eocene to Early Miocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 16pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Microchoeridae<sup>6<\/sup><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Necrolemur<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Long bony ear tubes. Tarsier-like lower limb adaptations for leaping.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Europe and Asia<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Early Eocene to Early Oligocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 1pt\">\n<td class=\"Table2-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\" colspan=\"4\">\n<p class=\"import-Normal\"><sup>1<\/sup> Derived from Fleagle 2013.<\/p>\n<p class=\"import-Normal\"><sup>2<\/sup> See Dunn Et al. 2016 and Rose Et al. 2018.<\/p>\n<p class=\"import-Normal\"><sup>3<\/sup> See Kirk and Williams 2011 and Seiffert Et al. 2009.<\/p>\n<p class=\"import-Normal\"><sup>4<\/sup> See Gregory 1920.<\/p>\n<p class=\"import-Normal\"><sup>5<\/sup> See Beard and MacPhee 1994 and Strait 2001.<\/p>\n<p class=\"import-Normal\"><sup>6<\/sup> See Schmid 1979.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\">\n<p class=\"import-Normal\">\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<p class=\"import-Normal\">Adapoids and omomyoids barely survived the Eocene-Oligocene extinctions, when colder temperatures, increased seasonality, and the retreat of rainforests to lower latitudes led to changes in mammalian biogeography. In North America, one genus (originally considered an omomyoid but recently reclassified as Adapoidea) persisted until the Miocene: <em>Ekgmowechashala<\/em> (Rose and Rensberger 1983). This taxon has highly unusual teeth and might have been a late immigrant to North America from Asia. In Asia, one family of adapoids, the Sivaladapidae, retained considerable diversity as late as the Late Miocene.<\/p>\n<h4 class=\"import-Normal\"><em>Adapoid Diversity<\/em><\/h4>\n<p class=\"import-Normal\">Adapoids were very diverse, particularly in the Eocene of North America and Europe. They can be divided into six families, with a few species of uncertain familial relationship. As a group, adapoids have some features in common, although much of what they share is plesiomorphic. Important features include the hallmarks of euprimates: postorbital bar, flattened nails, grasping extremities, and a petrosal bulla (Figures 9.9 and 9.10). In addition, some adapoids retain the ancestral dental formula of 2.1.4.3; that is, in each quadrant of the mouth, there are two incisors, one canine, four premolars, and three molars. In general, the incisors are small compared to the molars, but the canines are relatively large, with sexual dimorphism in some species. Cutting crests on the molars are well developed in some species, and the two halves of the mandible were fused at the midline in some species. Some adapoids were quite small (<em>Anchomomys <\/em>at a little over 100 g), and some were quite large (<em>Magnadapis<\/em> at 10 kg; 22 lbs.). Furthermore, the spaces and attachment features for the chewing muscles were truly enormous in some species, suggesting that these muscles were very large and powerful. Taken together, this suggests an overall adaptive profile of diurnal herbivory. The canine sexual dimorphism in some species suggests a possible mating pattern of polygyny, as males in polygynous primate species often compete with each other for mates and have especially large canine teeth.<\/p>\n<figure style=\"width: 548px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image18-1.jpg\" alt=\"Three partial animal crania.\" width=\"548\" height=\"350\" \/><figcaption class=\"wp-caption-text\">Figure 9.9: Representative crania of Adapidae from Museum d\u2019Histoire Naturelle Victor Brun, a natural history museum in Montauban, France. The white scale bar is 1 cm long. Credit: <a class=\"rId43\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Representative crania of adapids (European adapoids, (Figure 8.7)<\/a> from the <a class=\"rId44\" href=\"https:\/\/www.museum.montauban.com\/\">Museum d\u2019Histoire Naturelle Victor Brun in Montauban, France<\/a> original to <a class=\"rId45\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology <\/a>by Jonathan M. G. Perry is under a <a class=\"rId46\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<figure style=\"width: 547px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image19-2.jpg\" alt=\"Side views of small rodentlike skeleton with long tail.\" width=\"547\" height=\"525\" \/><figcaption class=\"wp-caption-text\">Figure 9.10: Darwinius masillae, a member of the Caenopithecidae. The slab on the left is Plate A and the slab on the right is Plate B. The parts of the skeleton in B that are outside of the dashed lines were fabricated. Credit: <a class=\"rId48\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Darwinius%20masillae%20holotype%20slabs.jpg\">Darwinius masillae holotype slabs<\/a> by Jens L. Franzen, Philip D. Gingerich, J\u00f6rg Habersetzer1, J\u00f8rn H. Hurum, Wighart von Koenigswald, B. Holly Smith is under a <a class=\"rId49\" href=\"https:\/\/creativecommons.org\/licenses\/by\/2.5\/legalcode\">CC BY 2.5 License<\/a>. Originally from Franzen Et al. 2009.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Omomyoid Diversity<\/em><\/h4>\n<p class=\"import-Normal\">Like adapoids, omomyoids appeared suddenly at the start of the Eocene and then became very diverse with most species dying out before the Oligocene. Omomyoids are known from thousands of jaws with teeth, relatively complete skulls for about a half-dozen species, and very little postcranial material. Omomyoids were relatively small primates, with the largest being less than three kilograms (approximately 7 lbs.; <em>Macrotarsius montanus<\/em>). All known crania possess a postorbital bar, which in some has been described as \u201cincipient closure.\u201d Some\u2014but not all\u2014known crania have an elongated bony ear tube extending lateral to the location of the eardrum, a feature seen in living tarsiers and <strong>catarrhines<\/strong>. The anterior teeth tend to be large, with canines that are usually not much larger than the incisors. Often it is difficult to distinguish closely related species using molar morphology, but the premolars tend to be distinct from one species to another. The postcranial skeleton of most omomyoids shows hallmarks of leaping behaviour reminiscent of that of tarsiers. In North America, omomyoids became very diverse and abundant. In fact, omomyoids from Wyoming are sufficiently abundant and from such stratigraphically controlled conditions that they have served as strong evidence for the gradual evolution of anatomical traits over time (Rose and Bown 1984).<\/p>\n<p class=\"import-Normal\"><em>Teilhardina <\/em>(Figure 9.11; see Figure 9.2) is one of the earliest and arguably the most plesiomorphic of omomyoids. <em>Teilhardina<\/em> has several species, most of which are from North America, with one from Europe (<em>T. belgica<\/em>) and one from Asia (<em>T. asiatica<\/em>). The species of this genus are anatomically similar and the deposits from which they are derived are roughly contemporaneous. Thus, this small primate likely dispersed across the northern continents very rapidly (Smith Et al. 2006).<\/p>\n<figure style=\"width: 545px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image3-1.jpg\" alt=\"World map with primates jumping across forested areas.\" width=\"545\" height=\"289\" \/><figcaption class=\"wp-caption-text\">Figure 9.11: A map of the world during the early Eocene showing one hypothesis for the direction of dispersal of the omomyoid Teilhardina. The map depicts primates hopping from continent to continent (East to West) via the forest corridors at high latitudes. Credit: <a href=\"https:\/\/www.pnas.org\/content\/103\/30\/11223\">Paleogeographic map showing hypothetical migration routes of Teilhardina (Figure 1)<\/a> by Thierry Smith, Kenneth D. Rose, and Philip D. Gingerich. 2006. <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">Proceedings of the National Academy of Sciences of the United States of America <\/a>103 (30): 11223\u201311227. Copyright (2006) National Academy of Sciences. Image <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">is used for non-commercial and educational purposes as outlined by PNAS.<\/a><\/figcaption><\/figure>\n<h2 class=\"import-Normal\">The Emergence of Modern Primate Groups<\/h2>\n<h3 class=\"import-Normal\"><strong>Origins of Crown Strepsirrhines<\/strong><\/h3>\n<p class=\"import-Normal\">Until the turn of this century, very little was known about the origins of the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1698\">crown<\/a><\/strong> (living) strepsirrhines. The Quaternary record of Madagascar contains many amazing forms of lemurs, including giant sloth-like lemurs, lemurs with perhaps monkey-like habits, lemurs with koala-like habits, and even a giant aye-aye (Godfrey and Jungers 2002). However, in Madagascar, early Tertiary continental sediments are lacking, and there is no record of lemur fossils before the Pleistocene.<\/p>\n<p class=\"import-Normal\">The fossil record of galagos is slightly more informative. Namely, there are Miocene African fossils that are very likely progenitors of lorisids (Simpson 1967). However, these are much like modern galagos and do not reveal anything about the relationship between crown strepsirrhines and Eocene fossil primates (but see below regarding <em>Propotto<\/em>). A similar situation exists for lorises in Asia: there are Miocene representatives, but these are substantially like modern lorises. The discovery of the first definite <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1699\"><strong>toothcomb<\/strong><\/a> canine (a hallmark of stresirrhines) in 2003 provided the \u201csmoking gun\u201d for the origin of crown strepsirrhines (Seiffert Et al. 2003). Recently, several other African primates have been recognized as having strepsirrhine affinities (Marivaux Et al. 2013; Seiffert 2012). The enigmatic Fayum primate <em>Plesiopithecus<\/em> is known from a skull that has been compared to aye-ayes and to lorises (Godinot 2006; Simons and Rasmussen 1994a).<\/p>\n<p class=\"import-Normal\">The now-recognized diversity of stem strepsirrhines from the Eocene and Oligocene of Afro-Arabia is strong evidence to suggest that strepsirrhines originated in that geographic area. This implies that lorises dispersed to Asia subsequent to an African origin. It is unknown what the first strepsirrhines in Madagascar were like. However, it seems likely that the lemuriform-lorisiform split occurred in continental Africa, followed by dispersal of lemuriform stock to Madagascar. Recent evidence suggests that <em>Propotto<\/em>, a Miocene primate from Kenya originally described as a potto antecedent, actually forms a clade with <em>Plesiopithecus <\/em>and the aye-aye; this might suggest that strepsirrhines dispersed to Madagascar from continental Africa more than once (Gunnell Et al. 2018).<\/p>\n<h3 class=\"import-Normal\"><strong>The Fossil Record of Tarsiers<\/strong><\/h3>\n<p class=\"import-Normal\">Tarsiers are so unusual that they fuel major debates about primate taxonomy. Tarsiers today are moderately diverse but geographically limited and not very different in their ecological habits\u2014especially considering that the split between them and their nearest living relative probably occurred over 50 million years ago. If omomyoids are excluded, then the fossil record of tarsiers is very limited. Two fossil species from the Miocene of Thailand have been placed in the genus <em>Tarsius<\/em>, as has an Eocene fossil from China (Beard Et al. 1994). These, and <em>Xanthorhysis<\/em> from the Eocene of China, are all very tarsier-like. In fact, it is striking that <em>Tarsius eocaenus<\/em> from China was already so tarsier-like as early as the Eocene. This suggests that tarsiers achieved their current morphology very early in their evolution and have remained more or less the same while other primates changed dramatically. Two additional genera, <em>Afrotarsius<\/em> from the Oligocene of Egypt and Libya and <em>Afrasia<\/em> from the Eocene of Myanmar, have also been implicated in tarsier origins, though the relationship between them and tarsiers is unclear (Chaimanee Et al. 2012). More recently, a partial skeleton of a small Eocene primate from China, <em>Archicebus achilles<\/em> (dated to approximately 55.8 million to 54.8 million years ago), was described as the most basal tarsiiform (Ni Et al. 2013). This primate is reconstructed as a diurnal insectivore and an arboreal quadruped that did some leaping\u2014but not to the specialized degree seen in living tarsiers. The anatomy of the eye in living tarsiers suggests that their lineage passed through a diurnal stage, so <em>Archicebus<\/em> (and diurnal omomyoids) might represent such a stage.<\/p>\n<h3 class=\"import-Normal\"><strong>Climate Change and the Paleogeography of Modern Primate Origins<\/strong><\/h3>\n<p class=\"import-Normal\">Changing global climate has had profound effects on primate dispersal patterns and ecological habits over evolutionary time. Primates today are strongly tied to patches of trees and particular plant parts such as fruits, seeds, and immature leaves. It is no surprise, then, that the distribution of primates mirrors the distribution of forests. Today, primates are most diverse in the tropics, especially in tropical rainforests. Global temperature trends across the Tertiary have affected primate ranges. Following the Cretaceous-Tertiary extinction event, cooler temperatures and greater seasonality characterized the Paleocene. In the Eocene, temperatures (and probably rainfall) increased globally and rainforests likely extended to very high latitudes. During this time, euprimates became diverse. With cooling and increased aridity at the end of the Eocene, many primate extinctions occurred in the northern continents and the surviving primates were confined to lower latitudes in South America, Afro-Arabia, Asia, and southern Europe. Among these survivors are the progenitors of the living groups of primates: lemurs and lorises, tarsiers, <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1700\"><strong>platyrrhines<\/strong><\/a> (monkeys of the Americas), and catarrhines (monkeys and apes of Africa and Asia) (Figure 9.12).<\/p>\n<figure id=\"attachment_277-4\" aria-describedby=\"caption-attachment-277-4\" style=\"width: 539px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" class=\"wp-image-264\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image2-5-e1691791570984.png\" alt=\"Map of world with gray continents.\" width=\"539\" height=\"306\" \/><figcaption id=\"caption-attachment-277-4\" class=\"wp-caption-text\">Figure 9.12: Map of key localities of early anthropoids on land that becomes Africa and southern Asia. Credit: <a class=\"rId56\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Oligocene Map with Key Early Anthropoid Localities (Figure 8.10)<\/a> original to <a class=\"rId57\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Elyssa Ebding at <a class=\"rId58\" href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\">GeoPlace, California State University, Chico<\/a> is under a <a class=\"rId59\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Localities based on Fleagle 2013, 265.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Competing Hypotheses for the Origin of Anthropoids<\/strong><\/h3>\n<p class=\"import-Normal\">There is considerable debate among paleoanthropologists as to the geographic origins of anthropoids. In addition, there is debate regarding the source group for anthropoids. Three different hypotheses have been articulated in the literature. These are the adapoid origin hypothesis, the omomyoid origin hypothesis, and the tarsier origin hypothesis (Figure 9.13).<\/p>\n<figure style=\"width: 419px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image24-1-1.jpg\" alt=\"Diagrams show three relationships among primate groups.\" width=\"419\" height=\"742\" \/><figcaption class=\"wp-caption-text\">Figure 9.13: Competing models of anthropoid origins. Branch lengths are not to scale. The omomyoid origin model and tarsier origin model do not make specific reference to the evolutionary position of strepsirrhines; however, they were included here for completeness. <a href=\"https:\/\/docs.google.com\/document\/d\/1VUDKMBJYS_jNONjLxT04jQN0_z9Ua50BRN6auGSHUuU\/edit\">A full text description of this image is available<\/a>. Credit: <a class=\"rId61\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Competing Trees for Anthropoid Origins (Figure 8.11)<\/a> original to <a class=\"rId62\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Jonathan M. G. Perry is under a <a class=\"rId63\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Adapoid Origin Hypothesis<\/em><\/h4>\n<p class=\"import-Normal\">Resemblances between some adapoids and some extant anthropoids include fusion of the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1702\"><strong>mandibular symphysis<\/strong><\/a>, overall robusticity of the chewing system, overall large body size, features that signal a diurnal lifestyle (like relatively small eye sockets), and ankle bone morphology. Another feature in common is canine sexual dimorphism, which is present in some species of adapoids (probably) and in several species of anthropoids.<\/p>\n<p class=\"import-Normal\">These features led some paleoanthropologists in the last half of the 20th century to suggest that anthropoids came from adapoid stock (Gingerich 1980; Simons and Rasmussen 1994b). One of the earliest supporters of the link between adapoids and anthropoids was Hans Georg Stehlin, who described much of the best material of adapoids and compared these Eocene primates to South American monkeys (Stehlin 1912). In more recent times, the adapoid origin hypothesis was reinforced by resemblances between these European adapoids (especially <em>Adapis <\/em>and <em>Leptadapis<\/em>) and some early anthropoids from the Fayum Basin (e.g., <em>Aegyptopithecus<\/em>, see below; Figure 9.14).<\/p>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 473.25pt\">\n<caption>Figure 9.14: Families of early anthropoids with example genera and traits: a table. Credit: Early anthropoids table original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Jonathan M. G. Perry and Stephanie L. Canington is under a <a class=\"rId64\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Content derived from Fleagle 2013.<\/caption>\n<thead>\n<tr style=\"height: 25pt\">\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Family<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Genera<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Morphology<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Location<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Age<\/strong><sup><strong>1<\/strong><\/sup><\/p>\n<p>&nbsp;<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table3-R\" style=\"height: 18pt\">\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Propliopithecidae<sup>2<\/sup><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Aegyptopithecus<\/em><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Large size. Cranial sexual dimorphism, large canines. Robust jaws and rounded molars. Partially ossified ear tube (in some). Robust skeleton; quadruped.<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Africa<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Late Eocene to Early Oligocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 16pt\">\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Parapithecidae<sup>3<\/sup><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Apidium<\/em><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Medium size. Retention of three premolars per quadrant. Rounded molars and premolars with large central cusps. Adaptations for leaping in the lower limb.<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Africa<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Late Eocene to Late Oligocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 16pt\">\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Proteopithecidae<sup>4<\/sup><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Proteopithecus<\/em><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Small size. Retention of three premolars per quadrant. Arboreal quadrupeds that ate fruit.<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Africa<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Late Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 16pt\">\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Oligopithecidae<sup>5<\/sup><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Catopithecus<\/em><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Small size. Skull has postorbital septum and unfused mandible. Deep jaws. Diet of fruits. Generalized quadruped.<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Africa<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Late Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 16pt\">\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Eosimiidae<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Eosimias<\/em><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Deep jaw with vertical unfused symphysis. Pointed incisors and canines. Crowded premolars.<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Asia<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Middle Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 16pt\">\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Amphipithecidae<sup>6<\/sup><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Pondaungia<\/em><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Deep jaws. Molars generally rounded with wide basins.<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Asia<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Middle Eocene to Early Oligocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 1pt\">\n<td class=\"Table3-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\" colspan=\"4\">\n<p class=\"import-Normal\"><sup>1<\/sup> Derived from Fleagle 2013.<\/p>\n<p class=\"import-Normal\"><sup>2<\/sup> See Gebo and Simons 1987 and Simons Et al. 2007.<\/p>\n<p class=\"import-Normal\"><sup>3<\/sup> See Feagle and Simons 1995 and Simons 2001.<\/p>\n<p class=\"import-Normal\"><sup>4<\/sup> See Simons and Seiffert 1999.<\/p>\n<p class=\"import-Normal\"><sup>5<\/sup> See Simons and Rasmussen 1996.<\/p>\n<p class=\"import-Normal\"><sup>6<\/sup> See Kay Et al. 2004.<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\">\n<p class=\"import-Normal\">\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<p class=\"import-Normal\">Unfortunately for the adapoid hypothesis, most of the shared features listed above probably emerged independently in the two groups as adaptations to a diet of hard and\/or tough foods. For example, fusion of the mandibular symphysis likely evolved as a means to strengthen the jaw against forces that would pull the two halves away from each other, in the context of active chewing muscles on both sides of the head generating great bite forces. This context would also favour the development of robust jaws, large chewing muscles, shorter faces, and some other features shared by some adapoids and some anthropoids.<\/p>\n<p class=\"import-Normal\">As older and more plesiomorphic anthropoids were found in the Fayum Basin, it became clear that the earliest anthropoids from Africa did not possess these features of jaw robusticity (Seiffert Et al. 2009). Furthermore, many adapoids never evolved these features. Fusion of the mandibular symphysis in adapoids is actually quite different from that in anthropoids and probably occurred during juvenile development in the former (Beecher 1983; Ravosa 1996). Eventually, the adapoid origin hypothesis fell out of favour among most paleoanthropologists, although the description of <em>Darwinius<\/em> is a recent revival of that idea (Franzen Et al. 2009; but see Seiffert Et al. 2009, Williams Et al. 2010b).<\/p>\n<h4 class=\"import-Normal\"><em>Omomyoid Origin Hypothesis<\/em><\/h4>\n<p class=\"import-Normal\">Similarities in cranial and hindlimb morphology between some omomyoids and extant tarsiers have led to the suggestion that tarsiers arose from some kind of omomyoid. In particular, <em>Necrolemur<\/em> has many features in common with tarsiers, as does the North American <em>Shoshonius<\/em>, which is known from a few beautifully preserved (although distorted) crania. Tarsiers and <em>Shoshonius <\/em>share exclusively some features of the base of the cranium; however, <em>Shoshonius<\/em> does not have any sign of postorbital closure, and it lacks the bony ear tube of tarsiers. Nevertheless, some of the resemblances between some omomyoids and tarsiers suggest that tarsiers might have originated from within the Omomyoidea (Beard 2002; Beard and MacPhee 1994). In this scenario, although living tarsiers and living anthropoids might be sister taxa, they might have evolved from different omomyoids, possibly separated from each other by more than 50 million years of evolution, or from anthropoids evolved from some non-omomyoid fossil group. The arguments against the omomyoid origin hypothesis are essentially the arguments <em>for<\/em> the tarsier origin hypothesis (see below). Namely, tarsiers and anthropoids share many features (especially of the soft tissues) that must have been retained for many millions of years or must have evolved convergently in the two groups. Furthermore, a key hard-tissue feature shared between the two extant groups, the postorbital septum, was not present in any omomyoid. Therefore, that feature must have arisen convergently in the two extant groups or must have been lost in omomyoids. Neither scenario is very appealing, although recent arguments for <strong>convergent evolution<\/strong> of the postorbital septum in tarsiers and anthropoids have arisen from embryology and histology of the structure (DeLeon Et al. 2016).<\/p>\n<h4 class=\"import-Normal\"><em>Tarsier Origin Hypothesis<\/em><\/h4>\n<p class=\"import-Normal\">Several paleoanthropologists have suggested that there is a relationship between tarsiers and anthropoids to the exclusion of omomyoids and adapoids (e.g., Cartmill and Kay 1978; Ross 2000; Williams and Kay 1995). Tarsiers and anthropoids today share several traits, including many soft-tissue features related to the olfactory system (e.g., the loss of a hairless external nose and loss of the median cleft running from the nose to the mouth, as possessed by strepsirrhines), and aspects of the visual system (e.g., the loss of a reflective layer at the back of the eye, similarities in carotid circulation to the brain, and mode of placentation). Unfortunately, none of these can be assessed directly in fossils. Some bony similarities between tarsiers and anthropoids include an extra air-filled chamber below the middle ear cavity, reduced bones within the nasal cavity, and substantial postorbital closure; these can be assessed in fossils, but the distribution of these traits in omomyoids does not yield clear answers. Furthermore, several similarities between tarsiers and anthropoids are probably due to similarities in sensory systems, which might have evolved in parallel for ecological reasons. Although early attempts to resolve the crown primates with molecular data were sometimes equivocal or in disagreement with one another, more recent analyses (including those of short interspersed elements) suggest that tarsiers and anthropoids are sister groups to the exclusion of lemurs and lorises (Williams Et al. 2010a). However, this does not address omomyoids, all of which are far too ancient for DNA extraction.<\/p>\n<p class=\"import-Normal\">The above three hypotheses are not the only possibilities for anthropoid origins. It may be that anthropoids are neither the closest sister group of tarsiers, nor evolved from adapoids or omomyoids. In recent years, two new groups of Eocene Asian primates have been implicated in the origin of anthropoids: the eosimiids and the amphipithecids. It is possible that one or the other of these two groups gave rise to anthropoids. Regardless of the true configuration of the tree for crown primates, the three major extant groups probably diverged from each other quite long ago (Seiffert Et al. 2004).<\/p>\n<h3 class=\"import-Normal\"><strong>Early Anthropoid Fossils in Africa<\/strong><\/h3>\n<figure style=\"width: 526px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image7-2.jpg\" alt=\"People digging in a sandy desert.\" width=\"526\" height=\"352\" \/><figcaption class=\"wp-caption-text\">Figure 9.15: Egyptian workers sweeping Quarry I in the Fayum Basin (2004). This technique, called wind harvesting, removes the desert crust and permits wind to blow out fine sediment and reveal fossils. Credit: <a class=\"rId66\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Egyptian workers sweeping Quarry I in the Fayum Basin (2004, Figure 8.12)<\/a> by Jonathan M. G. Perry is under a <a class=\"rId67\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<figure style=\"width: 280px\" class=\"wp-caption alignleft\"><img loading=\"lazy\" decoding=\"async\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image14-2.jpg\" alt=\"A person using a tool to expose bone in sand.\" width=\"280\" height=\"423\" \/><figcaption class=\"wp-caption-text\">Figure 9.16: Elwyn Laverne Simons excavating Aegyptopithecus in the Fayum Basin. Credit: <a class=\"rId69\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Elwyn Laverne Simons in the Fayum Basin (Figure 8.13)<\/a> used by permission of the <a class=\"rId70\" href=\"https:\/\/lemur.duke.edu\/\">Duke Lemur Center,<\/a> Division of Fossil Primates, is under a <a class=\"rId71\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">The classic localities yielding the greatest wealth of early anthropoid fossils are those from the Fayum Basin in Egypt (Simons 2008; Figure 9.15). The Fayum is a veritable oasis of fossil primates in an otherwise spotty early Tertiary African record. Since the 1960s, teams led by E. L. Simons have discovered several new species of early anthropoids, some of which are known from many parts of the skeleton and several individuals (Figure 9.16).<\/p>\n<p class=\"import-Normal\">The Fayum Jebel Qatrani Formation and Birket Qarun Formation between them have yielded a remarkable array of terrestrial, arboreal, and aquatic mammals. These include ungulates, bats, sea cows, elephants, hyraces, rodents, whales, and primates. Also, many other vertebrates, like water birds, were present. The area at the time of deposition (Late Eocene through Early Oligocene) was probably very wet, with slow-moving rivers, standing water, swampy conditions, and lots of trees (see Bown and Kraus 1988). In short, it was an excellent place for primates.<\/p>\n<h4 class=\"import-Normal\"><em>General Morphology of Anthropoids<\/em><\/h4>\n<p class=\"import-Normal\">The anthropoids known from the Fayum (and their close relatives from elsewhere in East Africa and Afro-Arabia) bear many of the anatomical hallmarks of extant anthropoids; however, there are plesiomorphic forms in several families that lack one or more anthropoid traits. All Fayum anthropoids known from skulls possess postorbital closure, most had fused mandibular symphyses, and most had ring-like <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1704\"><strong>ectotympanic<\/strong> <\/a> bones. Tooth formulae were generally either 2.1.3.3 or 2.1.2.3. Fayum anthropoids ranged in size from the very small <em>Qatrania<\/em> and <em>Biretia <\/em>(less than 500 g) to the much-larger <em>Aegyptopithecus<\/em> (approximately 7 kg; 15 lbs.). Fruit was probably the main component of the diet for most or all of the anthropoids, with some of them supplementing with leaves (Kay and Simons 1980; Kirk and Simons 2001; Teaford Et al. 1996). Most Fayum anthropoids were probably diurnal above-branch quadrupeds. Some of them (e.g., <em>Apidium<\/em>; see Figure 9.14) were probably very good leapers (Gebo and Simons 1987), but none show specializations for gibbon-style suspensory locomotion. Some of the Fayum anthropoids are known from hundreds of individuals, permitting the assessment of individual variation, sexual dimorphism, and in some cases growth and development. The description that follows provides greater detail for the two best known Fayum anthropoid families, the Propliopithecidae and the Parapithecidae; the additional families are summarized briefly.<\/p>\n<h4 class=\"import-Normal\"><em>Fayum Anthropoid Families<\/em><\/h4>\n<p class=\"import-Normal\">The Propliopithecidae (see Figure 9.14) include the largest anthropoids from the fauna, and they are known from several crania and some postcranial elements. They have been suggested to be stem catarrhines, although perhaps near the split between catarrhines and platyrrhines. The best known propliopithecid is <em>Aegyptopithecus<\/em>, known from many teeth, crania, and postcranial elements (Figure 9.17) .<\/p>\n<figure style=\"width: 431px\" class=\"wp-caption alignright\"><img loading=\"lazy\" decoding=\"async\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-2-1.jpg\" alt=\"Two animal skull side views.\" width=\"431\" height=\"281\" \/><figcaption class=\"wp-caption-text\">Figure 9.17: Female (left) and male (right) skull material for Aegyptopithecus zeuxis. The mandibles are not associated with the crania. Credit: <a href=\"https:\/\/www.pnas.org\/doi\/full\/10.1073\/pnas.0703129104#supplementary-materials\">Female and male cranium of A. zeuxi (03129Fig5, Supporting Information)<\/a> by Elwyn L. Simons, Erik R. Seiffert, Timothy M. Ryan, and Yousry Attia. 2007. <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">Proceedings of the National Academy of Sciences of the United States of America<\/a> 104 (21): 8731\u20138736. Copyright (2007) National Academy of Sciences. Image <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">is used for non-commercial and educational purposes as outlined by PNAS.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\">Parapithecidae are an extremely abundant and unusual family of anthropoids from the Fayum. The parapithecid <em>Apidium<\/em> is known from many jaws with teeth, crushed and distorted crania, and several skeletal elements. <em>Parapithecus<\/em> is known from cranial material including a beautiful, undistorted cranium. This genus shows extreme reduction of the incisors, including complete absence of the lower incisors in <em>P. grangeri <\/em>(Simons 2001). This trait is unique among primates. Parapithecids were once thought to be the ancestral stock of platyrrhines; however, their platyrrhine-like features are probably ancestral retentions, so the most conservative approach is to consider them stem anthropoids.<\/p>\n<p class=\"import-Normal\">The Proteopithecidae were small frugivores that probably mainly walked along horizontal branches on all fours. They are considered stem anthropoids. The best known genus, <em>Proteopithecus<\/em>, is represented by dentitions, crania, and post-cranial elements.<\/p>\n<p class=\"import-Normal\">The Oligopithecidae share a mixture of traits that makes them difficult to classify more specifically within anthropoids. The best known member, <em>Catopithecus<\/em>, is known from crania that demonstrate a postorbital septum and from mandibles that lack symphyseal fusion. They share the catarrhine tooth formula of 2.1.2.3 and have a canine honing complex that involves the anterior lower premolar. The postcranial elements known for the group suggest generalized arboreal quadrupedalism. The best known member, <em>Catopithecus<\/em>, is known from crania that demonstrate a postorbital septum and from mandibles that lack symphyseal fusion (Simons and Rasmussen 1996). The jaws are deep, with broad muscle attachment areas and crested teeth. <em>Catopithecus<\/em> was probably a little less than a kilogram in weight.<\/p>\n<p class=\"import-Normal\">Other genera of putative anthropoids from the Fayum include the very poorly known <em>Arsinoea<\/em>, the contentious <em>Afrotarsius<\/em>, and the enigmatic <em>Nosmips<\/em>. The last of these possesses traits of several major primate <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1705\">clades<\/a><\/strong> and defies classification (Seiffert Et al. 2010).<\/p>\n<h3 class=\"import-Normal\"><strong>Early Anthropoid Fossils in Asia<br style=\"clear: both\" \/><\/strong><\/h3>\n<p class=\"import-Normal\">For the last half of the 1900s, researchers believed that Africa was the unquestioned homeland of early anthropoids (see Fleagle and Kay 1994). However, two very different groups of primates from Asia soon began to change that. One was an entirely new discovery (Eosimiidae), and the other was a poorly known group discovered decades prior (Amphipithecidae). Soon, attention on anthropoid origins began to shift eastward (see Ross and Kay 2004; Simons 2004). If anthropoids arose in Asia instead of Africa, then this implies that the African early anthropoids either emigrated from Asia or evolved their anthropoid traits in parallel with living anthropoids.<\/p>\n<h4 class=\"import-Normal\"><em>Eosimiids<\/em><\/h4>\n<p class=\"import-Normal\">First described in the 1990s, the eosimiids are best represented by <em>Eosimias <\/em>(see Figure 9.14; Figure 9.18). This tiny \u201cdawn monkey\u201d is known from relatively complete jaws with teeth, a few small fragments of the face, and some postcranial elements (Beard Et al. 1994; Beard Et al. 1996; Gebo Et al. 2000). <em>Eosimias<\/em> (along with the other less-well-known genera in its family) bears some resemblance to tarsiers as well as anthropoids. Unfortunately, no good crania are known for this family, and the anatomy of, for example, the posterior orbital margin could be very revealing as to higher-level relationships.<\/p>\n<figure style=\"width: 550px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image16-1-1.jpg\" alt=\"Red-colored lower jaw of an animal.\" width=\"550\" height=\"232\" \/><figcaption class=\"wp-caption-text\">Figure 9.18: Cast of the right half of the mandible of Eosimias centennicus, type specimen. The white scale bar is 1 cm long. Credit: <a class=\"rId74\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Cast of the right half of the mandible of <\/a><a class=\"rId75\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\"><em>Eosimias centennicus <\/em><\/a><a class=\"rId76\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">(Figure 8.15),<\/a> type specimen, from K. D. Rose cast collection, photo by Jonathan M. G. Perry is under a <a class=\"rId77\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Amphipithecids<\/em><\/h4>\n<p class=\"import-Normal\">Amphipithecids are small- to medium-size primates (up to 10 kg; 22 lbs.). Most are from the Eocene Pondaung Formation in Myanmar (Early\u2013Middle Eocene), but one genus is known from Thailand. Some dental similarities with anthropoids were noted early on, such as deep jaws and wide basins that separate low molar cusps. The best known genera were <em>Pondaungia<\/em> and <em>Amphipithecus <\/em>(Ciochon and Gunnell 2002; see Figure 9.14). Another amphipithecid, <em>Siamopithecus<\/em> from Thailand, has very rounded molars and was probably a seed-eater (Figure 9.19). In addition to teeth and jaws, some cranial fragments, ankle material, and ends of post-cranial bones have been found for <em>Pondaungia<\/em>. There are important resemblances between these bones of <em>Pondaungia<\/em> and those of adapoids, suggesting adapoid affinities for the amphipithecidae. This would imply that the resemblances with anthropoids in the teeth are convergent, based on similarities in diet (see Ciochon and Gunnell 2002). Unfortunately, the association between post-cranial bones and teeth is not definite. With other primates in these faunas (including eosimiids), one cannot be certain that the post-cranial bones belong with the teeth. Some researchers suggest that some bones belong to a sivaladapid (or asiadapid) and others to an early anthropoid (Beard Et al. 2007; Marivaux Et al. 2003). Additional well-associated material of amphipithecids would help to clear up this uncertainty.<\/p>\n<figure style=\"width: 505px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image15-2.jpg\" alt=\"Four casts of jawbone fragments with teeth.\" width=\"505\" height=\"368\" \/><figcaption class=\"wp-caption-text\">Figure 9.19: Casts of representative amphipithecid material. A. Pondaungia cotteri right lower jaw fragment with m2 and m3. B. Siamopithecus eocaenus right upper jaw fragment with p4-m3. C. S. eocaenus right lower jaw fragment with partial m1, m2, and m3 in lateral view. D. Same as in C but occlusal view. White scale bars are 1 cm long. Credit: <a class=\"rId79\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Casts of representative amphipithecid material (Figure 8.16)l<\/a> from K. D. Rose cast collection, photo by Jonathan M. G. Perry is under a <a class=\"rId80\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Platyrrhine Dispersal to South America<\/strong><\/h3>\n<p class=\"import-Normal\">Today there is an impressive diversity of primates in South and Central America. These are considered to be part of a single clade, the Platyrrhini. Primates colonized South America sometime in the Eocene from an African source. In the first half of the 20th century, the source of platyrrhines was a matter of major debate among paleontologists, with some favouring a North American origin (e.g., Simpson 1940).<\/p>\n<p class=\"import-Normal\">Part of the reason for this debate is that South America was an island in the Eocene. Primates needed to cross open ocean to get there from either North America or Africa, although the distance from the former was shorter. Morphology yields clues to platyrrhine origins. The first known primates in South America have more in common morphologically with African primates than with North American ones. At the time, anthropoids were popping up in North Africa, whereas the only euprimates in North America were adapoids and omomyoids. Despite lacking a bony ear tube, early platyrrhines shared a great deal with other anthropoids, including full postorbital closure and fusion of the mandibular symphysis.<\/p>\n<p class=\"import-Normal\">The means by which a population of small North African primates managed to disperse across the Atlantic and survive to colonize South America remains a mystery. The most plausible scenario is one of rafting. That is, primates must have been trapped on vegetation that was blown out to sea by a storm. The vegetation then became a sort of life raft, which eventually landed ashore, dumping its passengers in South America. Rodents probably arrived in South America in the same way (Antoine Et al. 2012).<\/p>\n<p class=\"import-Normal\">Once ashore, platyrrhines must have crossed South America fairly rapidly because the earliest-known primates from that continent are from Peru (Bond Et al. 2015). Soon after that, platyrrhines were in Bolivia, namely <em>Branisella<\/em>. By the Miocene, platyrrhines were living in extreme southern Argentina and were exploiting a variety of feeding niches. The Early Miocene platyrrhines were all somewhat plesiomorphic in their morphology, but some features that likely arose by ecological convergence suggest (to some) relationships with extant platyrrhine families. This has led to a lively debate about the pattern of primate evolution in South America (Kay 2015; Kay &amp; Fleagle 2010; Rosenberger 2010). By the Middle Miocene, clear representatives of modern families were present in a diverse fauna from La Venta, Colombia (Wheeler 2010). The Plio-Pleistocene saw the emergence of giant platyrrhines as well as several taxa of platyrrhines living on Caribbean islands (Cooke Et al. 2016).<\/p>\n<p class=\"import-Normal\">The story of platyrrhines seems to be one of amazing sweepstakes dispersal, followed by rapid diversification and widespread geographic colonization of much of South America. After that, dramatic extinctions resulted in the current, much-smaller geographic distribution of platyrrhines. These extinctions were probably caused by changing climates, leading to the contraction of forests. Platyrrhines dispersed to the Caribbean and to Central America, with subsequent extinctions in those regions that might have been related to interactions with humans. Unlike anthropoids of Africa and Asia, platyrrhines do not seem to have evolved any primarily terrestrial forms and so have always been highly dependent on forests.<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Jonathan Perry and Primates of the Extreme South<\/h2>\n<p class=\"import-Normal\">Many primates are very vulnerable to ecological disturbance because they are heavily dependent on fruit to eat and trees to live in. This is one reason why so many primates are endangered today and why many of them went extinct due to climatic and vegetational changes in the past. I (Jonathan Perry) have conducted paleontological research focusing on primates that lived on the edge of their geographic distribution. This research has taken me to extreme environments in the Americas: southern Patagonia, the Canadian prairies, western Wyoming, and the badlands of eastern Oregon.<\/p>\n<p class=\"import-Normal\">Santa Cruz Province in Argentina is as far south as primates have ever lived. The Santa Cruz fauna of the Miocene has yielded a moderate diversity of platyrrhines, each with slightly different dietary adaptations. These include <em>Homunculus<\/em>, first described by Florentino Ameghino in 1891 (Figure 9.20). Recent fieldwork by my colleagues and I in Argentina has revealed several skulls of <em>Homunculus <\/em>as well as many parts of the skeleton (Kay Et al. 2012). The emerging profile of this extinct primate is one of a dedicated arboreal quadruped that fed on fruits and leaves. Many of the foods eaten by <em>Homunculus<\/em> must have been very tough and were probably covered and impregnated with grit; we suspect this because the cheek teeth are very worn down, even in young individuals, and because the molar tooth roots were very large, presumably to resist strong bite forces (Perry Et al. 2010, 2014).<\/p>\n<figure style=\"width: 497px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image9-2.jpg\" alt=\"An animal skull, a partial skull, and a fossil jaw with teeth.\" width=\"497\" height=\"634\" \/><figcaption class=\"wp-caption-text\">Figure 9.20: Representative specimens of Homunculus patagonicus. A. Adult cranium in lateral view. B. Adult cranium surface reconstructed from microCT scans, with the teeth segmented out. C. Juvenile cranium. White scale bars are 1cm long. Credit: <a class=\"rId82\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Representative specimens of <\/a><a class=\"rId83\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\"><em>Homunculus patagonicus <\/em><\/a><a class=\"rId84\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">(Figure 8.17)<\/a> photo by Jonathan M. G. Perry is under a <a class=\"rId85\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">I began working in Argentina while a graduate student at Duke University. I participated as a field assistant in a team led by my Ph.D. advisor, Richard F. Kay, and Argentine colleagues Sergio F. Vizca\u00edno and M. Susana Bargo. Most of the localities examined belong to a suite of beach sites known since the 1800s and visited by many field parties from various museums in the early 1900s. Since 2003, our international team of paleontologists from the U.S. and Argentina has visited these localities every single year (Figure 9.21). Over time, new fossils and new students have led to new projects and new approaches, including the use of microcomputed tomography (microCT) to visualize and analyse internal structures of the skeleton.<\/p>\n<figure style=\"width: 491px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image23.jpg\" alt=\"Sandy rocky coastline. People digging on a grassy hillside.\" width=\"491\" height=\"561\" \/><figcaption class=\"wp-caption-text\">Figure 9.21: Field localities in Argentina and Canada. A. Ca\u00f1adon Palos locality, coastal Santa Cruz Province, Argentina. B. Swift Current Creek locality, southwest Saskatchewan, Canada. Credits: A. <a class=\"rId87\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Ca\u00f1adon Palos Field Locality in Argentina<\/a> by Jonathan M. G. Perry is under a <a class=\"rId88\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. B. <a class=\"rId89\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Swift Current Creek locality, Saskatchewan, Canada<\/a> by Jonathan M. G. Perry is under a <a class=\"rId90\" 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\">Planet of Apes<\/h2>\n<h3 class=\"import-Normal\"><strong>Geologic Activity and Climate Change in the Miocene<\/strong><\/h3>\n<p class=\"import-Normal\">The Miocene Epoch was a time of mammalian diversification and extinction, global climate change, and ecological turnover. In the Miocene, there was an initial warming trend across the globe with the expansion of subtropical forests, followed by widespread cooling and drying with the retreat of tropical forests and replacement with more open woodlands and eventually grasslands. It was also a time of major geologic activity. On one side of the globe, South America experienced the rise of the Andes Mountains. On the other side, the Indian subcontinent collided with mainland Asia, resulting in the rise of the Himalayan Mountains. In Africa, volcanic activity promoted the development of the East African Rift System. Critical to the story of ape evolution was the exposure of an intercontinental landbridge between East Africa and Eurasia, permitting a true planet of apes (Figure 9.22).<\/p>\n<figure id=\"attachment_277-5\" aria-describedby=\"caption-attachment-277-5\" style=\"width: 580px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" class=\"wp-image-273\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image21-3-e1691792198797.png\" alt=\"Map of world with gray continents.\" width=\"580\" height=\"335\" \/><figcaption id=\"caption-attachment-277-5\" class=\"wp-caption-text\">Figure 9.22: Map of the world in the Miocene, highlighting fossil ape localities across Africa, southern Europe, and southern Asia. Credit: <a class=\"rId92\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Miocene Map with Fossil Ape Localities (Figure 8.19)<\/a> original to <a class=\"rId93\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Elyssa Ebding at <a class=\"rId94\" href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\">GeoPlace, California State University, Chico<\/a> is under a <a class=\"rId95\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Localities based on Fleagle 2013, 311.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Geographic Distribution: Africa, Asia, Europe<\/strong><\/h3>\n<p class=\"import-Normal\">The world of the Miocene had tremendous ape diversity compared to today. The earliest records of fossil apes are from Early Miocene deposits in Africa. However, something dramatic happened around 16 million years ago. With the closure of the ancient Tethys Sea, the subsequent exposure of the <em>Gomphotherium<\/em> Landbridge, and a period of global warming, the Middle\u2013Late Miocene saw waves of emigration of mammals (including primates) out of Africa and into Eurasia, with evidence of later African re-entry for some (Harrison 2010). Some of the mammals that dispersed from Africa to Eurasia and back were apes. Though most of these early apes left no modern descendants, some of them gave rise to the ancestors of modern apes\u2014including <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_800\">hominins<\/a><\/strong> (Figure 9.23).<\/p>\n<figure style=\"width: 560px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image20-1.jpg\" alt=\"Miocene apes set against a geologic time scale.\" width=\"560\" height=\"796\" \/><figcaption class=\"wp-caption-text\">Figure 9.23: Representative Miocene apes set against a geologic time scale. Credit: <a href=\"https:\/\/www.pnas.org\/content\/108\/14\/5554\">Range chart for Miocene hominoids of Western Eurasia (Figure 3)<\/a> by Isaac Casanovas-Vilar, David M. Alba, Miguel Garc\u00e9s, Josep M. Robles, and Salvador Moy\u00e0-Sol\u00e0. 2011. <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">Proceedings of the National Academy of Sciences of the United States of America<\/a> 108 (14): 5554-5559. Copyright (2011) National Academy of Sciences. Image <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">is used for non-commercial and educational purposes as outlined by PNAS.<\/a><\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Where Are the Monkeys? Diversity in the Miocene<\/strong><\/h3>\n<p class=\"import-Normal\">Whereas the Oligocene deposits in the Fayum of Egypt have yielded the earliest-known catarrhine fossils, the Miocene demonstrates some diversification of Cercopithecoidea. However, compared to the numerous and diverse Miocene apes (see below), monkeys of the Miocene are very rare and restricted to a single extinct family, the Victoriapithecidae (Figure 9.24). This family contains the earliest definite cercopithecoids.<\/p>\n<figure style=\"width: 357px\" class=\"wp-caption alignleft\"><img loading=\"lazy\" decoding=\"async\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image13-5.png\" alt=\"Front view of skull with pointed teeth.\" width=\"357\" height=\"322\" \/><figcaption class=\"wp-caption-text\">Figure 9.25: Skull of Victoriapithecus macinnesi. Credit: Victoriapithecus macinnesi skull photo taken at the Musee d&#8217;Histoire Naturelle, Paris by Ghedoghedo is under a CC BY-SA 3.0 License.<\/figcaption><\/figure>\n<p class=\"import-Normal\">These monkeys are found from northern and eastern Africa between 20 million and 12.5 million years ago (Miller Et al. 2009). The best known early African monkey is <em>Victoriapithecus <\/em>(Figure 9.25), a small-bodied (approximately 7 kg; 15 lbs.), small-brained monkey. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1708\">Bilophodonty<\/a><\/strong>, known to be a hallmark of molar teeth of modern cercopithecoid, was present to some extent in Victoriapithecids. <em>Victoriapithecus<\/em> has been reconstructed as being more frugivorous and perhaps spent more time on the ground (terrestrial locomotion) than in the trees (arboreal locomotion; Blue Et al. 2006). The two major groups of cercopithecoids today are cercopithecines and colobines. The earliest records demonstrating clear members of each of these two groups are at the end of the Miocene. Examples include the early colobine <em>Microcolobus<\/em> from Kenya and the early cercopithecine <em>Pliopapio<\/em> from Ethiopia.<\/p>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 473.25pt;height: 349px\">\n<caption>Figure 9.24: Some families of later anthropoids with example genera and traits: a table. Credit: Late anthropoids table original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Jonathan M. G. Perry and Stephanie L. Canington is under a <a class=\"rId100\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Content derived from Fleagle 2013.<\/caption>\n<thead>\n<tr style=\"height: 25pt\">\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 60px;width: 119.35px\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Family<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 60px;width: 103.417px\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Genera<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 60px;width: 191.65px\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Morphology<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 60px;width: 67.3667px\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Location<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 60px;width: 73.2167px\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Age<\/strong><sup><strong>1<\/strong><\/sup><\/p>\n<p>&nbsp;<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table4-R\" style=\"height: 18pt\">\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 77px;width: 119.35px\">\n<p class=\"import-Normal\">Victoriapithecidae<sup>2<\/sup><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 77px;width: 103.417px\">\n<p class=\"import-Normal\"><em>Victoriapithecus<\/em><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 77px;width: 191.65px\">\n<p class=\"import-Normal\">Long, sloping face. Round, narrowly spaced orbits. Deep cheek bones. Well-developed sagittal crest.<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 77px;width: 67.3667px\">\n<p class=\"import-Normal\">Africa<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 77px;width: 73.2167px\">\n<p class=\"import-Normal\">Early to Middle Miocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 16pt\">\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 61px;width: 119.35px\">\n<p class=\"import-Normal\">Proconsulidae<sup>3<\/sup><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 61px;width: 103.417px\">\n<p class=\"import-Normal\"><em>Proconsul<\/em><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 61px;width: 191.65px\">\n<p class=\"import-Normal\">Short face. Generalized dentition. Arboreal quadruped. Probably tailless.<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 61px;width: 67.3667px\">\n<p class=\"import-Normal\">Africa and Arabia<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 61px;width: 73.2167px\">\n<p class=\"import-Normal\">Early to Middle Miocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 16pt\">\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 46px;width: 119.35px\">\n<p class=\"import-Normal\">Pongidae<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 46px;width: 103.417px\">\n<p class=\"import-Normal\"><em>Gigantopithecus<\/em><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 46px;width: 191.65px\">\n<p class=\"import-Normal\">Largest primate ever. Deep jaws and low rounded molars.<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 46px;width: 67.3667px\">\n<p class=\"import-Normal\">Asia<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 46px;width: 73.2167px\">\n<p class=\"import-Normal\">Miocene to Present<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 1pt\">\n<td class=\"Table4-C\" style=\"border-color: #000000;border-style: solid none none;border-width: 0.5pt 0pt 0pt;padding: 0pt 5.4pt;height: 90px;width: 526.983px\" colspan=\"4\">\n<p class=\"import-Normal\"><sup>1<\/sup> Derived from Fleagle 2013.<\/p>\n<p class=\"import-Normal\"><sup>2<\/sup> See Benefit and McCrossin 1997 and Fleagle 2013.<\/p>\n<p class=\"import-Normal\"><sup>3<\/sup> See Begun 2007.<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"border-color: #000000;border-style: solid none none;border-width: 0.5pt 0pt 0pt;padding: 0pt 5.4pt;height: 90px;width: 73.2167px\">\n<p class=\"import-Normal\">\n<\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"height: 15px;width: 121.283px\"><\/td>\n<td style=\"height: 15px;width: 105.35px\"><\/td>\n<td style=\"height: 15px;width: 193.583px\"><\/td>\n<td style=\"height: 15px;width: 69.3px\"><\/td>\n<td style=\"height: 15px;width: 74.65px\"><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<h3><\/h3>\n<h3><\/h3>\n<h3><\/h3>\n<h3 class=\"import-Normal\"><strong>The Story of Us, the Apes<\/strong><\/h3>\n<h4 class=\"import-Normal\"><em>African Ape Diversity <\/em><\/h4>\n<p class=\"import-Normal\">The Early Miocene of Africa has yielded around 14 genera of early apes (Begun 2003). Many of these taxa have been reconstructed as frugivorous arboreal quadrupeds (Kay 1977). One of the best studied of these genera is the East African <em>Proconsul<\/em> (Family Proconsulidae; see Figure 9.24). Several species have been described, with body mass reconstructions ranging from 17 to 50 kg (approximately 37\u2013110 lbs.). A paleoenvironmental study reconstructed the habitat of <em>Proconsul <\/em>to be a dense, closed-canopy tropical forest (Michel Et al. 2014). No caudal vertebrae (tail bones) have been found in direct association with <em>Proconsul <\/em>post-crania, and the morphology of the sacrum is consistent with <em>Proconsul<\/em> lacking a tail (Russo 2016; Ward Et al. 1991).<\/p>\n<p class=\"import-Normal\">Overall, the African ape fossil record in the Late Miocene is sparse, with seven fossil localities dating between eleven and five million years ago (Pickford Et al. 2009). Nevertheless, most species of great apes live in Africa today. Where did the progenitors of modern African apes arise? Did they evolve in Africa or somewhere else? The paucity of apes in the Late Miocene of Africa stands in contrast to the situation in Eurasia. There, ape diversity was high. Furthermore, several Eurasian ape fossils show morphological affinities with modern hominoids (apes). Because of this, some paleoanthropologists suggest that the ancestors of modern African great apes recolonized Africa from Eurasia toward the end of the Miocene (Begun 2002). However, discoveries of Late Miocene hominoids like the Kenyan <em>Nakalipithecus<\/em> (9.9 million to 9.8 million years ago), the Ethiopian <em>Chororapithecus<\/em> (10.7 million to 10.1 million years ago), and the late-Middle Miocene Namibian <em>Otavipithecus<\/em> (13 million to 12 million years ago) fuel an alternative hypothesis\u2014namely that African hominoid diversity was maintained throughout the Miocene and that one of these taxa might, in fact, be the last common ancestor of extant African apes (Kunimatsu Et al. 2007; Mocke Et al. 2002). The previously underappreciated diversity of Late Miocene apes in Africa might be due to poor sampling of the fossil record in Africa.<\/p>\n<h4 class=\"import-Normal\"><em>Eurasian Ape Diversity<\/em><\/h4>\n<p class=\"import-Normal\">With the establishment of the <em>Gomphotherium<\/em> Landbridge (a result of the closure of the Eastern Mediterranean seaway; R\u00f6gl 1999), the Middle Miocene was an exciting time for hominoid radiations outside of Africa (see Figure 9.23). Eurasian hominoid species exploited their environments in many different ways in the Miocene. Food exploitation ranged from soft-fruit feeding in some taxa to hard-object feeding in others, in part owing to seasonal fluctuations and the necessary adoptions of fallback foods (DeMiguel Et al. 2014). For example, the molars of <em>Oreopithecus bambolii<\/em> (Family Hominidae) have relatively long lower-molar shearing crests, suggesting that this hominoid was very folivorous (Ungar and Kay 1995). Associated with variation in diet, there is great variation in the degree to which cranial features (e.g., zygomatic bone or supraorbital tori) are developed across the many taxa (Cameron 1997); however, Middle Miocene fossils tend to exhibit relatively thick molar enamel and relatively robust jaws (Andrews and Martin 1991).<\/p>\n<p class=\"import-Normal\">In Spain, the cranium with upper dentition, part of a mandible, and partial skeleton of <em>Pliobates <\/em>(Family Pliobatidae), a small-bodied ape (4\u20135 kg; 9\u201311 lbs.), was discovered in deposits dating to 11.6 million years ago (Alba Et al. 2015). It is believed to be a frugivore with a relative brain size that overlaps with modern cercopithecoids. The fossilized postcrania of <em>Pliobates<\/em> suggest that this ape might have had a unique style of locomotion, including the tendency to walk across the branches of trees with its palms facing downward and flexible wrists that permitted rotation of the forearm during climbing. However, the anatomy of the distal humerus differs from those of living apes in ways that suggest that <em>Pliobates<\/em> was less efficient at stabilizing its elbow while suspended (Benefit and McCrossin 2015). Two other recently described apes from Spain, <em>Pierolapithecus <\/em>and <em>Anoiapithecus<\/em>, are known from relatively complete skeletons. <em>Pierolapithecus<\/em> had a very projecting face and thick molar enamel as well as some skeletal features that suggest (albeit controversially) a less suspensory locomotor style than in extant apes (Moy\u00e0-Sol\u00e0 Et al. 2004). In contrast to <em>Pierolapithecus<\/em>, the slightly younger <em>Anoiapithecus<\/em> has a very flat face (Moy\u00e0-Sol\u00e0 Et al. 2009).<\/p>\n<p class=\"import-Normal\">Post-cranial evidence for suspensory or well-developed orthograde behaviours in apes does not appear until the Late Miocene of Europe. Primary evidence supporting these specialized locomotor modes includes the relatively short lumbar vertebrae of <em>Oreopithecus <\/em>(Figure 9.26) and <em>Dryopithecus<\/em> (Maclatchy 2004). Further, fossil material of the lower torso of <em>O. bambolii <\/em>(which dates to the <em>Pan<\/em>-hominin divergence) conveys a higher degree of flexion-extension abilities in the lumbar region (lower back) than what is possible in extant apes. Additionally, the hindlimb of <em>O. bambolii <\/em>is suggested to have supported powerful hip adduction during climbing (Hammond Et al. 2020). The Late Miocene saw the extinction of most of the Eurasian hominoids in an event referred to as the Vallesian Crisis (Agust\u00ed Et al. 2003). Among the latest surviving hominoid taxa in Eurasia were <em>Oreopithecus<\/em> and <em>Gigantopithecus<\/em>, the latter of which held out until the Pleistocene in Asia and was probably even sympatric with <em>Homo erectus<\/em> (Cachel 2015).<\/p>\n<figure style=\"width: 436px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image8-2-1.jpg\" alt=\"Posterior view of ancient ape skeleton.\" width=\"436\" height=\"775\" \/><figcaption class=\"wp-caption-text\">Figure 9.26: Skeleton of Oreopithecus bambolii. Credit: <a class=\"rId107\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Oreopithecus_bambolii_1.JPG\">Oreopithecus bambolii 1<\/a> by <a class=\"rId108\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Ghedoghedo\">Ghedoghedo<\/a> is under a <a class=\"rId109\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>The Origins of Extant Apes<\/strong><\/h3>\n<p class=\"import-Normal\">The fossil record of the extant apes is somewhat underwhelming: it ranges from being practically nonexistent for some taxa (e.g., chimpanzees) to being a little better for others (e.g., humans). There are many possible reasons for these differences in fossil abundance, and many are associated with the environmental conditions necessary for the fossilization of bones. One way to understand the evolution of extant apes that is not so dependent on the fossil record is via molecular evolutionary analyses. This can include counting up the differences in the genetic sequence between two closely related species to estimate the amount of time since these species shared a common ancestor. This is called a molecular clock, and it is often calibrated using fossils of known absolute age that stand in for the last common ancestor of a particular clade. Molecular clock estimates have placed the Hylobatidae and Hominidae split between 19.7 million and 24.1 million years ago, the African ape and Asian ape split between 15.7 million and 19.3 million years ago, and the split of Hylobatidae into its current genera between 6.4 million and 8 million years ago (Israfil Et al. 2011).<\/p>\n<h4 class=\"import-Normal\"><em>Small Ape Origins and Fossils<\/em><\/h4>\n<p class=\"import-Normal\">Unfortunately, the fossil record for the small (formerly \u201clesser\u201d) apes is meager, particularly in Miocene deposits. One possible early hylobatid is <em>Laccopithecus robustus<\/em>, a Late Miocene catarrhine from China (Harrison 2016). Although it does share some characteristics with modern gibbons and siamangs (including an overall small body size and a short face), <em>Laccopithecus<\/em> most likely represents a plesiomorphic stem catarrhine and is therefore distantly related to extant apes (Jablonski &amp; Chaplin 2009). A more likely candidate for the hylobatid stem is another Late Miocene taxon from China, <em>Yuanmoupithecus xiaoyuan<\/em>. Interpretation of its phylogenetic standing, however, is complicated by contradicting dental features\u2014some of them quite plesiomorphic\u2014which some believe best place <em>Yuanmoupithecus<\/em> as a stem hylobatid (Harrison 2016). Recently, a Middle Miocene Indian fossil ape, <em>Kapi ramnagarensis<\/em>, has extended the fossil record of small apes by approximately five million years. Its teeth are suggestive of a shift to a more frugivorous diet and it is likely a stem hylobatid (Gilbert Et al. 2020). The history of Hylobatidae becomes clearer in the Pleistocene, with fossils representing extant genera.<\/p>\n<h4 class=\"import-Normal\"><em>Great Ape Origins and Fossils<\/em><\/h4>\n<p class=\"import-Normal\">The most extensive fossil record of a modern great ape is that of our own genus, <em>Homo<\/em>. However, the evolutionary history of the Asian great ape, the orangutan (<em>Pongo<\/em>), is becoming clearer. Today, orangutans are found only on the islands of Borneo and Sumatra. However, Pleistocene-aged teeth, attributed to <em>Pongo<\/em>, have been found in Cambodia, China, Laos, Peninsular Malaysia, and Vietnam\u2014demonstrating the vastness of the orangutan\u2019s previous range (Ibrahim Et al. 2013; Wang Et al. 2014). <em>Sivapithecus <\/em>from the Miocene of India and Pakistan is represented by many specimens, including parts of the face. <em>Sivapithecus<\/em> is very similar to <em>Pongo<\/em>, especially in the face, and it probably is closely related to ancestral orangutans (Pilbeam 1982). Originally, jaws and teeth belonging to the former genus <em>Ramapithecus<\/em> were thought to be important in the origin of humans (Simons 1961), but now these are recognized as specimens of <em>Sivapithecus<\/em> (Kelley 2002). Postcranial bones of <em>Sivapithecus<\/em>, however, suggest a more generalized locomotor mode\u2014including terrestrial locomotion\u2014than seen in <em>Pongo <\/em>(Pilbeam Et al. 1990). Stable carbon and oxygen isotope data from dental enamel have reconstructed the paleoecological space of <em>Sivapithecus <\/em>(as well as the contemporaneous Late Miocene pongine <em>Khoratpithecus<\/em>) within the canopies of forested habitats (Habinger Et al. 2022).<\/p>\n<p class=\"import-Normal\">A probable close relative of <em>Sivapithecus <\/em>is the amazing <em>Gigantopithecus<\/em> (see Figure 9.24). Known only from teeth and jaws from China and India (e.g., Figure 9.27), this ape probably weighed as much as 270 kg (595 lbs.) and was likely the largest primate ever (Bocherens Et al. 2017). Because of unique features of its teeth (including molarized premolars and patterns of wear) and its massive size, it has been reconstructed as a bamboo specialist, somewhat like the modern panda. Small silica particles (phytoliths) from grasses have been found stuck to the molars of <em>Gigantopithecus<\/em> (Ciochon Et al. 1990). Recent studies evaluating the carbon isotope composition of the enamel sampled from <em>Gigantopithecus<\/em> teeth suggest that this ape exploited a wide range of vegetation, including fruits, leaves, roots, and bamboo (Bocherens Et al. 2017). Its face is reminiscent of that of modern orangutans and it might belong in the same family, Pongidae (Kelley 2002).<\/p>\n<figure style=\"width: 488px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image12.jpg\" alt=\"Superior view of mandible and teeth.\" width=\"488\" height=\"533\" \/><figcaption class=\"wp-caption-text\">Figure 9.27: Cast of the mandible of Gigantopithecus blacki. Credit: <a class=\"rId111\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Gigantopithecus%20blacki%20mandible%20010112.jpg\">Gigantopithecus blacki mandible 010112<\/a> by <a class=\"rId112\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Wilson44691\">Wilson44691<\/a> is under a <a class=\"rId113\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">In Africa, the first fossil to be confidently attributed to <em>Pan<\/em>, and known to be the earliest evidence of a chimpanzee, was described based on teeth found in Middle Pleistocene deposits in the Eastern Rift Valley of Kenya (McBrearty and Jablonski 2005). Paleoenvironmental reconstructions of this locality suggest that this early chimpanzee was living in close proximity to early <em>Homo<\/em> in a closed-canopy wooded habitat. Similarly, fossil teeth and mandibular remains attributed to two species of Middle-Late Miocene apes\u2014<em>Chororapithecus abyssinicus<\/em> (from Ethiopia; Suwa Et al. 2007) and <em>Nakalipithecus nakayamai<\/em> (from Kenya; Kunimatsu Et al. 2007)\u2014have been suggested as basal members of the gorilla clade.<\/p>\n<p class=\"import-Normal\">While the deposits of Eastern Africa have yielded a profound record of our fossil hominin ancestors, the continent\u2019s rainforests remain a \u201cpalaeontological desert\u201d (Rosas Et al. 2022). Clearly, more work is needed to fill in the large gaps in the fossil record of the nonhuman great apes. The twentieth century witnessed the discovery of many hominin fossils in East Africa, which have been critical for improving our understanding of human evolution. While twenty-first-century conservationists fight to prevent the extinction of the living great apes, perhaps efforts by twenty-first-century paleoanthropologists will yield the evolutionary story of these, our closest relatives.<\/p>\n<h2>Cultural Connections and Contemporary Implications<\/h2>\n<p>Even today, continued study of primate evolution and the gathering of new evidence are critical to our understanding of our biological history. The Piltdown Man hoax is a striking example of how, without modern carbon dating technology and continued study in the field, false conclusions and evidence can be accepted as truth. In 1912, Charles Dawson\u2014an amateur antiquarian and archeologist\u2014presented a combination of human skull fragments and an ape-like lower jaw as evidence of a previously unknown \u2018missing link\u2019 in the line of human ancestors (Groote et al., 2016). Immediately, members of the British archaeological establishment accepted the specimen because it seemed consistent with the belief that a larger\u2014human-like\u2014brain was among the earliest features to evolve in humans. This specimen remained largely accepted for decades until, in 1953, modern chemical analyses proved that the remains had been deliberately altered: the jaw had been stained, the teeth filed, and the skull fragments were subsequently carbon-dated to only approximately 600 years old (Australian Museum, n.d.).<\/p>\n<p>Among many such cases, the Piltdown Man hoax demonstrates why the continued study of primate evolution remains imperative to contemporary biological anthropology. As new discoveries, technologies, and analytical methods provide additional evidence, our understanding of human biological history can be continually evaluated and refined.<\/p>\n<p>Discussion questions<\/p>\n<ul>\n<li style=\"font-weight: 400\">Can you think of an example where a belief that was widely accepted as scientific truth became harmful, not only to science but also to the people within that society?<\/li>\n<li style=\"font-weight: 400\">Why is it important for scientists to remain open to revising their conclusions when new evidence emerges? How does the Piltdown Man hoax demonstrate the importance of this?<\/li>\n<\/ul>\n<div class=\"textbox shaded\">\n<h2>Summary<\/h2>\n<p>While there are large gaps in the fossil records linking primates to early hominins, evolutionary trends make it clear that humans are one branch of the broader primate family tree. In this chapter we go over the major development that characterize primate evolution: enhanced vision, grasping hands and feet, greater reliance on social behaviour, and increased brain complexity. It is these traits which distinguish primates from other mammals and furthermore, help define major subsections within the primate class, such as strepsirrhines, haplorhines, monkeys, apes, and ultimately hominins.<\/p>\n<p>Within this chapter, we also examine how anthropologists reconstruct these evolutionary relationships. Fossil evidence has provided key information about when and where different primates lived, while genetic data such as skeletal features allow researchers to understand how extinct species moved, ate, and interacted with their environments. Just as crucial are the influences of Earth\u2019s changing environments: continental drift, glacial cycles, and long-term climate shifts have repeatedly reshaped habitats, driving both extinctions and the emergence of new adaptive forms, including the emergence of our own human lineage.<\/p>\n<h2 class=\"import-Normal\">Review Questions<strong><br \/>\n<\/strong><\/h2>\n<ul>\n<li>Compare three major hypotheses about primate origins, making reference to each one\u2019s key ecological reason for primate uniqueness.<\/li>\n<li>Explain how changes in temperature, rainfall, and vegetation led to major changes in primate biogeography over the Early Tertiary.<\/li>\n<li>List some euprimate features that plesiadapiforms have and some that they lack.<\/li>\n<li>Contrast adapoids and omomyoids in terms of life habits.<\/li>\n<li>Describe one piece of evidence for each of the adapoid, omomyoid, and tarsier origin hypotheses for anthropoids.<\/li>\n<li>Discuss the biogeography of the origins of African great apes and orangutans using examples from the Miocene ape fossil record.<\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\">Key Terms<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\"><strong>Adapoidea<\/strong>: Order: Primates. One of the earliest groups of euprimates (true primates; earliest records from the early Eocene).<\/p>\n<p class=\"import-Normal\"><strong>A<\/strong><strong>daptive radiations<\/strong>: Rapid diversifications of single lineages into many species which may present unique morphological features in response to different ecological settings.<\/p>\n<p class=\"import-Normal\"><strong>Ancestral traits<\/strong>: Features that were inherited from a common ancestor and which remain (largely) unchanged.<\/p>\n<p class=\"import-Normal\"><strong>Anthropoids<\/strong>:Group containing monkeys and apes, including humans.<\/p>\n<p class=\"import-Normal\"><strong>Auditory bulla<\/strong>: The rounded bony floor of the middle ear cavity.<\/p>\n<p class=\"import-Normal\"><strong>Bilophodonty<\/strong>: Dental condition in which the cusps of molar teeth form ridges (or lophs) separated from each other by valleys (seen, e.g., in modern catarrhine monkeys).<\/p>\n<p class=\"import-Normal\"><strong>Catarrhines<\/strong>: Order: Primates; Suborder: Anthropoidea; Infraorder: Catarrhini. Group, with origins in Africa and Asia, that contains monkeys and apes, including humans.<\/p>\n<p class=\"import-Normal\"><strong>Clade<\/strong>:Group containing all of the descendants of a single ancestor. A portion of a phylogenetic tree represented as a bifurcation (node) in a lineage and all of the branches leading forward in time from that bifurcation.<\/p>\n<p class=\"import-Normal\"><strong>Convergent evolution<\/strong>: The independent evolution of a morphological feature in animals not closely related (e.g., wings in birds and bats).<\/p>\n<p class=\"import-Normal\"><strong>Crown<\/strong>: Smallest monophyletic group (clade) containing a specified set of extant taxa and all descendants of their last common ancestor.<\/p>\n<p class=\"import-Normal\"><strong>Diastema<\/strong>: Space between adjacent teeth.<\/p>\n<p class=\"import-Normal\"><strong>Diffuse coevolution<\/strong>: The ecological interaction between whole groups of species (e.g., primates) with whole groups of other species (e.g., fruiting trees).<\/p>\n<p class=\"import-Normal\"><strong>Ectotympanic<\/strong>: Bony ring or tube that holds the tympanic membrane (eardrum).<\/p>\n<p class=\"import-Normal\"><strong>Euprimates<\/strong>: Order: Primates. True primates or primates of modern aspect.<\/p>\n<p class=\"import-Normal\"><strong>Haplorhines<\/strong>: Group containing catarrhines, platyrrhines, and tarsiers.<\/p>\n<p class=\"import-Normal\"><strong>Hominins<\/strong>: Modern humans and any extinct relatives more closely related to us than to chimpanzees.<\/p>\n<p class=\"import-Normal\"><strong>Mandibular symphysis<\/strong>: Fibrocartilaginous joint between the left and right mandibular segments, located in the midline of the body.<\/p>\n<p class=\"import-Normal\"><strong>Omomyoidea<\/strong>: Order: Primates; Superfamily: Omomyoidea. One of the earliest groups of euprimates (true primates; earliest record in the early Eocene).<\/p>\n<p class=\"import-Normal\"><strong>Petrosal bone<\/strong>: The portion of the temporal bone that houses the inner ear apparatus.<\/p>\n<p class=\"import-Normal\"><strong>Plagiaulacoid<\/strong>: Dental condition where at least one of the lower cheek-teeth (molars or premolars) is a laterally compressed blade.<\/p>\n<p class=\"import-Normal\"><strong>Platyrrhines<\/strong>: Order: Primates; Suborder: Anthropoidea; Infraorder: Platyrrhini. Group containing monkeys found in the Americas.<\/p>\n<p class=\"import-Normal\"><strong>Plesiadapiforms<\/strong>: Order: Plesiadapiformes. Archaic primates or primate-like placental mammals (Early Paleocene\u2013Late Eocene).<\/p>\n<p class=\"import-Normal\"><strong>P<\/strong><strong>lesiomorphic<\/strong>: Having features that are shared by different groups which arose from a common ancestor.<\/p>\n<p class=\"import-Normal\"><strong>Stem<\/strong>: Taxa that are basal to a given crown group but are more closely related to the crown group than to the closest living sister taxon of the crown group.<\/p>\n<p class=\"import-Normal\"><strong>Strepsirrhines<\/strong>: Order: Primates; Suborder: Stresirrhini. Group containing lemurs, lorises, and galagos (does not include tarsiers).<\/p>\n<p class=\"import-Normal\"><strong>Toothcomb<\/strong>: Dental condition found in modern strepsirrhines in which the lower incisors and canines are laterally compressed and protrude forward at a nearly horizontal inclination. This structure is used in grooming.<\/p>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<h2 class=\"import-Normal\">For Further Exploration<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\">Beard, Chris. 2004. <em>The Hunt for the Dawn Monkey: Unearthing the Origins of Monkeys, Apes, and Humans<\/em>. Berkeley: University of California Press.<\/p>\n<p class=\"import-Normal\">Begun, David R. 2010. \u201cMiocene Hominids and the Origins of the African Apes and Humans.\u201d <em>Annual Review of Anthropology<\/em> 39: 67\u201384.<\/p>\n<p class=\"import-Normal\">Fleagle, John G. 2013. <em>Primate Adaptation and Evolution.<\/em> Third edition. San Diego, CA: Academic Press.<\/p>\n<p class=\"import-Normal\">Gebo, Daniel L., ed. 1993. <em>Postcranial Adaptations in Nonhuman Primates<\/em>. Dekalb: Northern Illinois University Press.<\/p>\n<p class=\"import-Normal\">Godfrey, Laurie R., and William L. Jungers. 2002. \u201cQuaternary Fossil Lemurs.\u201d In <em>The Primate Fossil Record, <\/em>edited by Walter C. Hartwig, 97\u2013121. Cambridge: Cambridge University Press.<\/p>\n<p class=\"import-Normal\">Godinot, Marc. 2006. \u201cLemuriform Origins as Viewed from the Fossil Record.\u201d <em>Folia Primatologica<\/em> 77 (6): 446\u2013464.<\/p>\n<p class=\"import-Normal\">Kay, Richard F. 2018. \u201c100 Years of Primate Paleontology.\u201d <em>American Journal of Physical Anthropology<\/em> 165 (4): 652\u2013676.<\/p>\n<p class=\"import-Normal\">Marivaux, Laurent. 2006. \u201cThe Eosimiid and Amphipithecid Primates (Anthropoidea) from the Oligocene of the Bugti Hills (Balochistan, Pakistan): New Insight into Early Higher Primate Evolution in South Asia.\u201d <em>Palaeovertebrata, Montpellier <\/em>34 (1\u20132): 29\u2013109.<\/p>\n<p class=\"import-Normal\">Martin, R. D. 1990. <em>Primate Origins and Evolution<\/em><em>: A <\/em><em>Phylogenetic Reconstruction<\/em>. Princeton: Princeton University Press.<\/p>\n<p class=\"import-Normal\">Rose, Kenneth D., Marc Godinot, and Thomas M. Bown. 1994. \u201cThe Early Radiation of Euprimates and the Initial Diversification of Omomyidae.\u201d In <em>Anthropoid Origins: The Fossil Evidence, <\/em>edited by John G. Fleagle and Richard F. Kay, 1\u201328. New York: Plenum Press.<\/p>\n<p class=\"import-Normal\">Ross, Callum F. 1999. \u201cHow to Carry Out Functional Morphology.\u201d <em>Evolutionary Anthropology<\/em> 7 (6): 217\u2013222.<\/p>\n<p class=\"import-Normal\">Seiffert, Erik R. 2012. \u201cEarly Primate Evolution in Afro-Arabia.\u201d Evolutionary Anthropology: Issues, News, and Reviews 21(6): 239\u2013253.<\/p>\n<p class=\"import-Normal\">Szalay, Frederic S., and Eric Delson. 1979. Evolutionary History of the Primates. New York: Academic Press.<\/p>\n<p class=\"import-Normal\">Ungar, Peter S. 2002. \u201cReconstructing the Diets of Fossil Primates.\u201d In <em>Reconstructing Behavior in the Primate Fossil Record<\/em>, edited by Joseph Plavcan, Richard F. Kay, William Jungers, and Carel P. van Schaik, 261\u2013296. New York: Kluwer Academic\/Plenum Publishers.<\/p>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\">Agust\u00ed, J., A. Sanz de Siria, and M. Garc\u00e9s M. 2003. \u201cExplaining the End of the Hominoid Experiment in Europe.\u201d <em>Journal of Human Evolution<\/em> 45 (2): 145\u2013153.<\/p>\n<p class=\"import-Normal\">Alba, David M., Sergio Alm\u00e9cija, Daniel DeMiguel, Josep Fortuny, Miriam P\u00e9rez de los R\u00edos, Marta Pina, Josep M. Robles, and Salvador Moy\u00e0-Sol\u00e0. 2015. \u201cMiocene Small-Bodied Ape from Eurasia Sheds Light on Hominoid Evolution.\u201d <em>Science<\/em> 350 (6260): aab2625.<\/p>\n<p class=\"import-Normal\">Andrews, Peter, and Lawrence Martin. 1991. \u201cHominoid Dietary Evolution.\u201d <em>Philosophical Transactions of the Royal Society of London B: Biological Sciences<\/em> 334 (1270): 199\u2013209.<\/p>\n<p class=\"import-Normal\">Antoine, Pierre-Oliver, Laurent Marivaux, Darren A. Croft, Guillaume Billet, Morgan Ganer\u00f8d, Carlos Jaramillo, Thomas Martin, et al. 2012. \u201cMiddle Eocene Rodents from Peruvian Amazonia Reveal the Pattern and Timing of Caviomorph Origins and Biogeography.\u201d <em>Proceedings of the Royal Society B: Biological Sciences<\/em> 279 (1732): 1319\u20131326.<\/p>\n<p class=\"import-Normal\">Beard, K. 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Christopher, Laurent Marivaux, Soe Thura Tun, Aung Naing Soe, Yaowalak Chaimanee, Wanna Htoon, Bernard Marandat, Htun Htun Aung, and Jean-Jacques Jaeger. 2007. \u201cNew Sivaladapid Primates from the Eocene Pondaung Formation of Myanmar and the Anthropoid Status of Amphipithecidae.\u201d <em>Bulletin of Carnegie Museum of Natural History<\/em> 39: 67\u201376.<\/p>\n<p class=\"import-Normal\">Beard, K. Christopher, Tao Qi, Mary R. Dawson, Banyue Wang, and Chuankuei Li. 1994. \u201cA Diverse New Primate Fauna from Middle Eocene Fissure-Fillings in Southeastern China.\u201d <em>Nature<\/em> 368 (6472): 604\u2013609.<\/p>\n<p class=\"import-Normal\">Beard, K. Christopher, Yongsheng Tong, Mary R. 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Christopher Beard, Aung Aung Kyaw, Aung Naing Soe, Chit Sein, Vincent Lazzari, et al. 2012. \u201cLate Middle Eocene Primate from Myanmar and the Initial Anthropoid Colonization of Africa.\u201d <em>Proceedings of the National Academy of Sciences<\/em> <em>of the United States of America <\/em>109 (26): 10293\u201310297.<\/p>\n<p class=\"import-Normal\">Chester, Stephen G. B., Jonathan I. Bloch, Doug M. Boyer, and William A. Clemens. 2015. \u201cOldest Known Euarchontan Tarsals and Affinities of Paleocene <em>Purgatorius<\/em> to Primates.\u201d <em>Proceedings of the National Academy of Sciences of the United States of America<\/em> 112 (5): 1487\u20131492.<\/p>\n<p class=\"import-Normal\">Ciochon, Russell L., and Gregg F. Gunnell. 2002. \u201cChronology of Primate Discoveries in Myanmar: Influences on the Anthropoid Origins Debate.\u201d <em>Yearbook of Physical Anthropology<\/em> 45(S35): 2\u201335.<\/p>\n<p class=\"import-Normal\">Ciochon, R. L., D. R. Piperno, and R. G. Thompson. 1990. \u201cOpal Phytoliths Found on the Teeth of the Extinct Ape <em>Gigantopithecus blacki<\/em>: Implications for Paleodietary Studies.\u201d <em>Proceedings of the National Academy of Sciences of the United States of America<\/em> 87 (20): 8120\u20138124.<\/p>\n<p class=\"import-Normal\">Clemens, William A. 2004. \u201c<em>Purgatorius<\/em> (Plesiadapiformes, Primates?, Mammalia), a Paleocene Immigrant into Northeastern Montana: Stratigraphic Occurrences and Incisor Proportions.\u201d <em>Bulletin of Carnegie Museum of Natural History<\/em> 36: 3\u201313.<\/p>\n<p class=\"import-Normal\">Cooke, Siobh\u00e1n B., Justin T. Gladman, Lauren B. Halenar, Zachary S. Klukkert, and Alfred L. 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Christopher Beard, Tao Qi, and Jingwen Wang. 2000. \u201cThe Oldest Known Anthropoid Postcranial Fossils and the Early Evolution of Higher Primates.\u201d <em>Nature<\/em> 404 (6775): 276\u2013278.<\/p>\n<p class=\"import-Normal\">Gebo, Daniel L., and Elwyn L. Simons. 1987. \u201cMorphology and Locomotor Adaptations of the Foot in Early Oligocene Anthropoids.\u201d <em>American Journal of Physical Anthropology<\/em> 74 (1): 83\u2013101.<\/p>\n<p class=\"import-Normal\">Gilbert, Christopher C., Alejandra Ortiz, Kelsey D. Pugh, Christopher J. Campisano, Biren A. Patel, Ningthoujam Premjit Singh, John G. Fleagle, and Rajeev Patnaik. 2020. \u201cNew Middle Miocene Ape (Primates: Hylobatidae) from Ramnagar, India, Fills Major Gaps in the Hominoid Fossil Record.\u201d <em>Proceedings of the Royal Society B<\/em> 287(1934): 20201655.<\/p>\n<p class=\"import-Normal\">Gingerich, P. 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Renne. 2021. \u201cEarliest Palaeocene Purgatoriids and the Initial Radiation of Stem Primates.\u201d <em>Royal Society Open Science<\/em> 8(2):210050. doi:10.1098\/rsos.210050.<\/p>\n<h2 class=\"import-Normal\">Acknowledgements<\/h2>\n<p class=\"import-Normal\">We are immensely grateful to the editors of this book, Drs. Beth Shook, Lara Braff, Katie Nelson, and Kelsie Aguilera, for their time and commitment to making this knowledge freely accessible to all, and for giving us the opportunity to participate in this important project.<\/p>\n<\/div>\n<\/div>\n<div class=\"glossary\"><span class=\"screen-reader-text\" id=\"definition\">definition<\/span><template id=\"term_1683_1683\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1683\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<div class=\"textbox\">\n<p>Learn more about <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/about-the-authors\/\">authors and editors<\/a>,<a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/student-contributors\/\"> Hess' student contributions<\/a>, and <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/versioning-history\/\">versioning history<\/a>.<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\">Learning Objectives<\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li>Understand the major trends in primate evolution from the origin of primates to the origin of our own species.<\/li>\n<li>Learn about primate adaptations and how they characterize major primate groups.<\/li>\n<li>Discuss the kinds of evidence that anthropologists use to find out how extinct primates are related to each other and to living primates.<\/li>\n<li>Recognize how the changing geography and climate of Earth have influenced where and when primates have thrived or gone extinct.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p class=\"import-Normal\">The first fifty million years of primate evolution was a series of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1683\">adaptive radiations<\/a><\/strong> leading to the diversification of the earliest lemurs, monkeys, and apes. The primate story begins in the canopy and understory of conifer-dominated forests, with our small, furtive ancestors subsisting at night, beneath the notice of day-active dinosaurs.<\/p>\n<p class=\"import-Normal\">From the ancient <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1684\"><strong>plesiadapiforms<\/strong><\/a> (archaic primates) to the earliest groups of true primates (<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1686\"><strong>euprimates<\/strong><\/a>) (Bloch and Boyer 2002), the origin of our own order is characterized by the struggle for new food sources and microhabitats in the arboreal setting. Climate change forced major extinctions as the northern continents became increasingly dry, cold, and seasonal and as tropical rainforests gave way to deciduous forests, woodlands, and eventually grasslands. Lemurs, lorises, and tarsiers\u2014once diverse groups containing many species\u2014became rare, except for lemurs in Madagascar, where there were no anthropoid competitors and perhaps few predators. Meanwhile, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1685\">anthropoids<\/a><\/strong> (monkeys and apes) likely emerged in Asia and then dispersed across parts of the northern hemisphere, Africa, and ultimately South America. The movement of continents, shifting sea levels, and changing patterns of rainfall and vegetation contributed to the developing landscape of primate biogeography, morphology, and behaviour. Today\u2019s primates provide modest reminders of the past diversity and remarkable adaptations of their extinct relatives. This chapter explores the major trends in primate evolution from the origin of the Order Primates to the beginnings of our own lineage, providing a window into these stories from our ancient past.<\/p>\n<h2 class=\"import-Normal\">Major Hypotheses About Primate Origins<\/h2>\n<p class=\"import-Normal\">For many groups of mammals, there is a key feature that led to their success. A good example is powered flight in bats. Primates lack a feature like this (see Chapter 6). Instead, if there is something unique about primates, it is probably a group of features rather than one single thing. Because of this, anthropologists and palaeontologists struggle to describe an ecological scenario that could explain the rise and success of our own order. Three major hypotheses have been advanced to consider the origin of primates and to explain what makes our order distinct among mammals (Figure 9.1); these are described below.<\/p>\n<figure id=\"attachment_277-6\" aria-describedby=\"caption-attachment-277-6\" style=\"width: 634px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-255\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2023\/03\/8.1.jpg\" alt=\"Primates swinging in tree, eating an insect, and eating fruit.\" width=\"634\" height=\"221\" \/><figcaption id=\"caption-attachment-277-6\" class=\"wp-caption-text\">Figure 9.1: The three major hypotheses are (a) the arboreal hypothesis, (b) the visual predation hypothesis, and (c) the angiosperm-primate coevolution hypothesis. Credit: Primate origin hypotheses original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by <a class=\"rId13\" href=\"https:\/\/marynelsonstudio.com\">Mary Nelson<\/a> is under a <a class=\"rId14\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Arboreal Hypothesis<\/strong><\/h3>\n<p class=\"import-Normal\">In the 1800s, many anthropologists viewed all animals in relation to humans. That is, animals that were more like humans were considered to be more \u201cadvanced\u201d and those lacking humanlike features were considered more \u201cprimitive.\u201d This way of thinking was particularly obvious in studies of primates. A more modern way of referring to members of a group that lack certain evolutionary innovations seen in other members is to call them <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1688\"><strong>plesiomorphic<\/strong><\/a> (literally \u201canciently shaped\u201d). The state of their morphological features is sometimes referred to as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1689\"><strong>ancestral<\/strong><strong> traits<\/strong><\/a>.<\/p>\n<p class=\"import-Normal\">Thus, when anthropologists sought features that separate primates from other mammals, they focused on features that were least developed in lemurs and lorises, more developed in monkeys, and most developed in apes (Figure 9.2). Frederic Wood Jones, one of the leading anatomist-anthropologists of the early 1900s, is usually credited with the Arboreal Hypothesis of primate origins (Jones 1916). This hypothesis holds that many of the features of primates evolved to improve locomotion in the trees; this way of getting around is referred to as arboreal. For example, the grasping hands and feet of primates are well suited to gripping tree branches of various sizes and our flexible joints are good for reorienting the extremities in many different ways. A mentor of Jones, Grafton Elliot Smith, had suggested that the reduced olfactory system, acute vision, and forward-facing eyes of primates are adaptations for making accurate leaps and bounds through a complex, three-dimensional canopy (Smith 1912). The forward orientation of the eyes in primates causes the visual fields to overlap, enhancing depth perception, especially at close range. Evidence to support this hypothesis includes the facts that many extant primates are arboreal, and the plesiomorphic members of most primate groups are dedicated arborealists. The Arboreal Hypothesis was well accepted by most anthropologists at the time and for decades afterward.<\/p>\n<figure style=\"width: 663px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image27-2.png\" alt=\"Diagram shows primates descended from Plesiadapiforms.\" width=\"663\" height=\"543\" \/><figcaption class=\"wp-caption-text\">Figure 9.2: Primate family tree showing major groups. Disconnected lines show uncertainty about relationships. Two lines lead to tarsiers from different possible groups of origin. <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 class=\"rId16\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Primate family tree (Figure 8.2)<\/a> by Jonathan M. G. Perry 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<h3 class=\"import-Normal\"><strong>Visual Predation Hypothesis<\/strong><\/h3>\n<p class=\"import-Normal\">In the late 1960s and early 1970s, Matt Cartmill studied and tested the idea that the characteristic features of primates evolved in the context of arboreal locomotion. Cartmill noted that squirrels climb trees (and even vertical walls) very effectively, even though they lack some of the key adaptations of primates. As members of the Order Rodentia, squirrels also lack the hand and foot anatomy of primates. They have claws instead of flattened nails and their eyes face more laterally than those of primates. Cartmill reasoned that there must be some other explanation for the unique traits of primates. He noted that some nonarboreal animals share at least some of these traits with primates; for example, cats and predatory birds have forward-facing eyes that enable visual field overlap. Cartmill suggested that the unique suite of features in primates is an adaptation to detecting insect prey and guiding the hands (or feet) to catch insects (Cartmill 1972). His hypothesis emphasizes the primary role of vision in prey detection and capture; it is explicitly comparative, relying on form-function relationships in other mammals and nonmammalian vertebrates. According to Cartmill, many of the key features of primates evolved for preying on insects in this special manner (Cartmill 1974).<\/p>\n<h3 class=\"import-Normal\"><strong>Angiosperm-Primate Coevolution Hypothesis<\/strong><\/h3>\n<p class=\"import-Normal\">The visual predation hypothesis was unpopular with some anthropologists. One reason for this is that many primates today are not especially predatory. Another is that, whereas primates do seem well adapted to moving around in the smallest, terminal branches of trees, insects are not necessarily easier to find there. A counterargument to the visual predation hypothesis is the angiosperm-primate coevolution hypothesis. Primate ecologist Robert Sussman (Sussman 1991) argued that the few primates that eat mostly insects often catch their prey on the ground rather than in tree branches. Furthermore, predatory primates often use their ears more than their eyes to detect prey. Finally, most early primate fossils show signs of having been omnivorous rather than insectivorous. Instead, he argued, the earliest primates were probably seeking fruit. Fruit (and flowers) of angiosperms (flowering plants) often develop in the terminal branches. Therefore, any mammal trying to access those fruits must possess anatomical traits that allow them to maintain their hold on thin branches and avoid falling while reaching for the fruits. Primates likely evolved their distinctive visual traits and extremities in the Paleocene (approximately 65 million to 54 million years ago) and Eocene (approximately 54 million to 34 million years ago) epochs, just when angiosperms were going through a revolution of their own\u2014the evolution of large, fleshy fruit that would have been attractive to a small arboreal mammal. Sussman argued that, just as primates were evolving anatomical traits that made them more efficient fruit foragers, angiosperms were also evolving fruit that would be more attractive to primates to promote better seed dispersal. This mutually beneficial relationship between the angiosperms and the primates was termed coevolution or more specifically <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1691\"><strong>diffuse coevolution<\/strong>.<\/a><\/p>\n<p class=\"import-Normal\">At about the same time, D. Tab Rasmussen noted several parallel traits in primates and the South American woolly opossum, <em>Caluromys<\/em>. He argued that early primates were probably foraging on both fruits and insects (Rasmussen 1990). As is true of <em>Caluromys<\/em> today, early primates probably foraged for fruits in the terminal branches of angiosperms, and they probably used their visual sense to aid in catching insects. Insects are also attracted to fruit (and flowers), so these insects represent a convenient opportunity for a primarily fruit-eating primate to gather protein. This solution is a compromise between the visual predation hypothesis and the angiosperm-primate coevolution hypothesis. It is worth noting that other models of primate origins have been proposed, and these include the possibility that no single ecological scenario can account for the origin of primates.<\/p>\n<h2 class=\"import-Normal\">The Origins of Primates<\/h2>\n<h3 class=\"import-Normal\"><strong>Paleocene: Mammals in the Wake of Dinosaur Extinctions<\/strong><\/h3>\n<p class=\"import-Normal\">Placental mammals, including primates, originated in the Mesozoic Era (approximately 251 million to 65.5 million years ago), the Age of Dinosaurs. During this time, most placental mammals were small, probably nocturnal, and probably avoided predators via camouflage and slow, quiet movement. It has been suggested that the success and diversity of the dinosaurs constituted a kind of ecological barrier to Mesozoic mammals. The extinction of the dinosaurs (and many other organisms) at the end of the Cretaceous Period (approximately 145.5\u201365.5 million years ago) might have opened up these ecological niches, leading to the increased diversity and disparity in mammals of the Tertiary Period (approximately 65.5\u20132.5 million years ago).<\/p>\n<p class=\"import-Normal\">The Paleocene was the first epoch in the Age of Mammals. Soon after the Cretaceous-Tertiary (K-T) extinction event, new groups of placental mammals appear in the fossil record. Many of these groups achieved a broad range of sizes and lifestyles as well as a great number of species before declining sometime in the Eocene (or soon thereafter). These groups were ultimately replaced by the modern orders of placental mammals (Figure 9.3). It is unknown whether these replacements occurred gradually, for example by competitive exclusion, or rapidly, perhaps by sudden geographic dispersals with replacement. In some senses, the Paleocene might have been a time of recovery from the extinction event; it was cooler and more seasonal globally than the subsequent Eocene.<\/p>\n<figure style=\"width: 628px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image26.jpg\" alt=\"Person in front of a mural depicting forest animals.\" width=\"628\" height=\"511\" \/><figcaption class=\"wp-caption-text\">Figure 9.3: A mural of Eocene flora and fauna in North America. Credit: <a class=\"rId19\" href=\"https:\/\/flickr.com\/photos\/126377022@N07\/18404106406\">Image from page 27 of \"Annual report for the year ended June 30 ...\" (1951)<\/a> by <a class=\"rId20\" href=\"https:\/\/flickr.com\/photos\/internetarchivebookimages\/\">Internet Archive Book Images<\/a> has been designated to the <a class=\"rId21\" href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/\">public domain (CC0)<\/a>. This photograph of the mural \"Fauna and flora of middle Eocene in the Wyoming area\" by Jay Matternes, was originally published by the <a class=\"rId22\" href=\"https:\/\/www.si.edu\/\">Smithsonian<\/a>, and can be viewed in context in the <a class=\"rId23\" href=\"https:\/\/archive.org\/details\/annualreportfory1961united\/page\/7\/mode\/1up?view=theater\">online version of this book<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Plesiadapiforms, the Archaic Primates<\/strong><\/h3>\n<p class=\"import-Normal\">The Paleocene epoch saw the emergence of several families of mammals that have been implicated in the origin of primates. These are the plesiadapiforms, which are archaic primates, meaning they possessed some primate features and lacked others. The word <em>plesiadapiform <\/em>means \u201calmost adapiform,\u201d a reference to some similarities between some plesiadapiforms and some adapiforms (or adapoids; later-appearing true primates)\u2014mainly in the molar teeth. Because enamel fossilizes better than other parts of the body, the molar teeth are the parts most often found and first discovered for any new species. Thus, dental similarities were often the first to be noticed by early mammalian paleontologists, partly explaining why plesiadapiforms were thought to be primates. Major morphological differences between plesidapiforms and euprimates (true primates) were observed later when more parts of plesiadapiform skeletons were discovered. Many plesiadapiforms have unusual anterior teeth and most have digits possessing claws rather than nails. So far, no plesiadapiform ever discovered has a postorbital bar (seen in extant <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1712\">strepsirrhines<\/a><\/strong>) or septum (as seen in <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1713\">haplorhines<\/a><\/strong>), and whether or not the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1711\">auditory bulla<\/a><\/strong> was formed by the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1714\"><strong>petrosal bone<\/strong> <\/a>remains unclear for many plesiadapiform specimens. Nevertheless, there are compelling reasons (partly from new skeletal material) for including plesiadapiforms within the Order Primates.<\/p>\n<h4 class=\"import-Normal\"><em>Geographic and Temporal Distribution<\/em><\/h4>\n<p class=\"import-Normal\"><em>Purgatorius<\/em> is generally considered to be the earliest primate. This Paleocene mammal is known from teeth that are very plesiomorphic for a primate. It has some characteristics that suggest it is a basal plesiadapiform, but there is very little to link it specifically with euprimates (see Clemens 2004). Its ankle bones suggest a high degree of mobility, signaling an arboreal lifestyle (Chester Et al. 2015). <em>Purgatorius<\/em> is plesiomorphic enough to have given rise to all primates, including the plesiadapiforms. However, new finds suggest that this genus was more diverse and had more differing tooth morphologies than previously appreciated (Wilson Mantilla Et al. 2021). Plesiadapiform families were numerous and diverse during parts of the Paleocene in western North America and western Europe, with some genera (e.g., <em>Plesiadapis<\/em>; see Figure 9.4) living on both continents (Figure 9.5). Thus, there were probably corridors for plesiadapiform dispersal between the two continents, and it stands to reason that these mammals were living all across North America, including in the eastern half of the continent and at high latitudes. A few plesiadapiforms have been described from Asia (e.g., <em>Carpocristes<\/em>), but the affinities of these remain uncertain.<\/p>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 473.25pt\">\n<caption>Figure 9.4: Families of plesiadapiforms with example genera and traits: a table. Credit: Plesiadapiforms table original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Jonathan M. G. Perry and Stephanie L. Canington is under a <a class=\"rId24\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Content derived from Fleagle 2013.<\/caption>\n<thead>\n<tr style=\"height: 25pt\">\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Family<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Genera<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Morphology<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Location<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Age<\/strong><sup><strong>1<\/strong><\/sup><\/p>\n<p>&nbsp;<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table1-R\" style=\"height: 17pt\">\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Paromomyidae<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Ignacius<\/em><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Long, dagger-like, lower incisor.<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">North America and Europe<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Early Paleocene to Late Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 18pt\">\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Carpolestidae<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Carpolestes<\/em><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Plagiaulacoid dentition. Limb adaptations to terminal branch feeding. Grasping big toe.<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">North America, Europe, and Asia<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Middle Paleocene to Early Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 16pt\">\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Plesiadapidae<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Plesiadapis<\/em><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Mitten-like upper incisor. Diastema. Large body size for group.<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">North America and Europe<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Middle Paleocene to Early Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 1pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\" colspan=\"4\">\n<p class=\"import-Normal\"><sup>1<\/sup> Derived from Fleagle 2013.<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\">\n<p class=\"import-Normal\">\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<figure id=\"attachment_277-7\" aria-describedby=\"caption-attachment-277-7\" style=\"width: 555px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-258 \" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5-5-e1691791897574.png\" alt=\"Global map with not fully formed continents.\" width=\"555\" height=\"308\" \/><figcaption id=\"caption-attachment-277-7\" class=\"wp-caption-text\">Figure 9.5: Map of the world in the Paleocene, highlighting plesiadapiform localities on lands that would become North America, southern Europe, and eastern Asia. Credit: <a class=\"rId26\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Paleocene Map with Plesiadapiform Localities (Figure 8.4)<\/a> original to<a class=\"rId27\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"> Expl<\/a><a class=\"rId28\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">orations: An Open Invitation to Biological Anthropology<\/a> by Elyssa Ebding at <a class=\"rId29\" href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\">GeoPlace, California State University, Chico<\/a> is under a <a class=\"rId30\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Localities based on Fleagle 2013, 211.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>General Morphological Features<\/em><\/h4>\n<p class=\"import-Normal\">Although there is much morphological variation among the families of plesiadapiforms, some common features unite the group. Most plesiadapiforms were small, the largest being about three kilograms (approximately 7 lbs.; <em>Plesiadapis cookei<\/em>). They had small brains and fairly large snouts, with eyes that faced more laterally than in euprimates. Many species show reduction and\/or loss of the canine and anterior premolars, with the resulting formation of a rodent-like <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1172\">diastema<\/a> <\/strong>(a pronounced gap between the premolars and the incisors, with loss of at least the canine); this probably implies a herbivorous diet. Some families appear to have had very specialized diets, as suggested by unusual tooth and jaw shapes.<\/p>\n<p class=\"import-Normal\">Arguably the most interesting and unusual family of plesiadapiforms is the Carpolestidae. They are almost exclusively from North America (with a couple of possible members from Asia), and mainly from the Middle and Late Paleocene. Their molars are not very remarkable, being quite similar to those of some other plesiadapiforms (e.g., Plesiadapidae). However, their lower posterior premolars (p4) are laterally compressed and blade-like with vertical serrations topped by tiny cuspules. This unusual dental morphology is termed <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1693\"><strong><em>plagiaulacoid<\/em><\/strong> <\/a> (Simpson 1933). The upper premolar occlusal surfaces are broad and are covered with many small cuspules; the blade-like lower premolar might have cut across these cuspules, between them, or both.<\/p>\n<div class=\"mceTemp\"><\/div>\n<figure style=\"width: 314px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image1-5.png\" alt=\"Small brown animal with long tail.\" width=\"314\" height=\"178\" \/><figcaption class=\"wp-caption-text\">Figure 9.6: An artistic rendition of Carpolestes simpsoni moving along a small diameter support. Credit: <a class=\"rId32\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:CarpolestesCL.png\">CarpolestesCL<\/a> by <a class=\"rId33\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Sisyphos23\">Sisyphos23<\/a> is under a <a class=\"rId34\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Many plesiadapiforms have robust limb bones with hallmarks of arboreality. Instead of having nails, most taxa had sharp claws on most or all of the digits. The extremities show grasping abilities comparable to those of primates and some arboreal marsupials. Nearly complete skeletons have yielded a tremendous wealth of information on locomotor and foraging habits. Many plesiadapiforms appear to have been able to cling to vertical substrates (like a broad tree trunk) using their sharp claws, propelling themselves upward using powerful hindlimbs, bounding along horizontal supports, grasping smaller branches, and moving head-first down tree trunks. In carpolestids in particular, the skeleton appears to have been especially well adapted to moving slowly and carefully in small terminal branches (Figure 9.6).<\/p>\n<\/div>\n<div class=\"textbox shaded\">\n<h3 class=\"import-Normal\">Dig Deeper: Debate on the Relationship of Plesiadapiforms to True Primates<\/h3>\n<p class=\"import-Normal\">In the middle of the twentieth century, treeshrews (Order Scandentia) were often considered part of the Order Primates, based on anatomical similarities between some treeshrews and primates. For many people, plesiadapiforms represented intermediates between primates and treeshrews, so plesiadapiforms were included in Primates as well.<\/p>\n<p class=\"import-Normal\">Studies of reproduction and brain anatomy in treeshrews and lemurs suggested that treeshrews are not primates (e.g., Martin 1968). This was soon followed by the suggestion to also expel plesiadapiforms (Martin 1972) from the Order Primates. Like treeshrews, plesiadapiforms lack a postorbital bar, nails, and details of the ear region that characterize true primates. Many paleoanthropologists were reluctant to accept this move to banish plesiadapiforms (e.g., F. S. Szalay, P. D. Gingerich).<\/p>\n<p class=\"import-Normal\">Later, K. Christopher Beard (1990) found that in some ways, the digits of paromomyid plesiadapiforms are actually more similar to those of dermopterans (Order Dermoptera), the closest living relatives of primates, than they are to those of primates themselves (but see Krause 1991). At the same time, Richard Kay and colleagues (1990) found that cranial circulation patterns and auditory bulla morphology in the paromomyid, <em>Ignacius <\/em>(see Figure 9.4), are more like those of dermopterans than of primates.<\/p>\n<p class=\"import-Normal\">For many anthropologists, this one-two punch effectively removed plesiadapiforms from the Order Primates. In the last two decades, the tide of opinion has turned again, with many researchers reinstating plesiadapiforms as members of the Order Primates. New and more complete specimens demonstrate that the postcranial skeletons of plesiadapiforms, including the hands and feet, were primate-like, not dermorpteran-like (Bloch and Boyer 2002, 2007). New fine-grained CT scans of relatively complete plesiadapiform skulls revealed that they share some key traits with primates to the exclusion of other placental mammals (Bloch and Silcox 2006). Most significant was the suggestion that <em>Carpolestes simpsoni <\/em>possessed an auditory bulla formed by the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1696\"><strong>petrosal <\/strong><strong>bone<\/strong><\/a>, like in all living primates.<\/p>\n<p class=\"import-Normal\">The debate about the status of plesiadapiforms continues, owing to a persistent lack of key bones in some species and owing to genuine complexity of the anatomical traits involved. Maybe plesiadapiforms were the ancestral stock from which all primates arose, with some plesiadapiforms (e.g., carpolestids) nearer to the primate <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1723\">stem<\/a><\/strong> than others.<\/p>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<h3 class=\"import-Normal\"><strong>Adapoids and Omomyoids, the First True Primates<\/strong><\/h3>\n<h4 class=\"import-Normal\"><em>Geographic and Temporal Distribution<\/em><\/h4>\n<p class=\"import-Normal\">The first universally accepted fossil primates are the adapoids (Superfamily <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1695\"><strong>Adapoidea<\/strong><\/a>) and the omomyoids (Superfamily <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1694\">Omomyoidea<\/a>)<\/strong>. These groups become quite distinct over evolutionary time, filling mutually exclusive niches for the most part. However, the earliest adapoids are very similar to the earliest omomyoids.<\/p>\n<p class=\"import-Normal\">The adapoids were mainly diurnal and herbivorous, with some achieving larger sizes than any plesiadapiforms (10 kg; 22 lbs.). By contrast, the omomyoids were mainly nocturnal, insectivorous and frugivorous, and small.<\/p>\n<p class=\"import-Normal\">Both groups appear suddenly at the start of the Eocene, where they are present in western North America, western Europe, and India (Figure 9.7). This wide dispersal of early primates was probably due to the presence of rainforest corridors extending far into northern latitudes.<\/p>\n<figure id=\"attachment_277-8\" aria-describedby=\"caption-attachment-277-8\" style=\"width: 539px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-260\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image22-3-e1691792023503.png\" alt=\"Global map with not fully formed continents and omomyoid localities.\" width=\"539\" height=\"317\" \/><figcaption id=\"caption-attachment-277-8\" class=\"wp-caption-text\">Figure 9.7: Map of the world in the Eocene, highlighting adapoid and omomyoid localities on lands that would become North America, southern Europe, Africa, and Asia. Credit: <a class=\"rId36\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Eocene Map with Adapoid and Omomyoid Localities (Figure 8.6)<\/a> original to <a class=\"rId37\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Elyssa Ebding at <a class=\"rId38\" href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\">GeoPlace, California State University, Chico<\/a> is under a <a class=\"rId39\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Localities based on Fleagle 2013, 229.<\/figcaption><\/figure>\n<p class=\"import-Normal\">In North America and Europe, both groups achieved considerable diversity in the Middle Eocene, then mostly died out at the end of that epoch (Figure 9.8). In some Eocene rock formations in the western United States, adapoids and omomyoids make up a major part of the mammalian fauna. The Eocene of India has yielded a modest diversity of euprimates, some of which are so plesiomorphic that it is difficult to know whether they are adapoids or omomyoids (or even early anthropoids).<\/p>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 473.25pt\">\n<caption>Figure 9.8: Families of adapoids and omomyoids with example genera and traits: a table. Credit: Adapoids and omomyoids table original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Jonathan M. G. Perry and Stephanie L. Canington is under a <a class=\"rId40\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Content derived from Fleagle 2013.<\/caption>\n<thead>\n<tr style=\"height: 25pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Family<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Genera<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Morphology<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Location<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Age<\/strong><sup><strong>1<\/strong><\/sup><\/p>\n<p>&nbsp;<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table2-R\" style=\"height: 18pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Cercamoniidae<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Donrussellia<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Variable in tooth number and jaw shape.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Europe and Asia<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Early to Late Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 16pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Asiadapidae<sup>2<\/sup><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Asiadapis<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Plesiomorphic teeth and jaw resemble early Omomyids.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Asia<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Early Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 16pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Caenopithecidae<sup>3<\/sup><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Darwinius<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Robust jaws with crested molars. Fewer premolars.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Europe, Africa, North America, and Asia<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Middle to Late Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 16pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Adapidae<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Adapis<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Fused mandible. Long molar crests. Large size and large chewing muscles.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Europe<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Late Eocene to Early Oligocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 16pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Sivaladapidae<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Sivaladapis<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Some large with robust jaws.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Asia<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Middle Eocene to Late Miocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 16pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Notharctidae<sup>4<\/sup><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Notharctus<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Canine sexual dimorphism. Lemur-like skull. Clinging and leaping adaptations.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">North America and Europe<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Early to Middle Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 16pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Omomyidae<sup>5<\/sup><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Teilhardina<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Small, nocturnal, frugivorous or insectivorous. Tarsier-like skull in some.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">North America, Europe, and Asia<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Early Eocene to Early Miocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 16pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Microchoeridae<sup>6<\/sup><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Necrolemur<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Long bony ear tubes. Tarsier-like lower limb adaptations for leaping.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Europe and Asia<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Early Eocene to Early Oligocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 1pt\">\n<td class=\"Table2-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\" colspan=\"4\">\n<p class=\"import-Normal\"><sup>1<\/sup> Derived from Fleagle 2013.<\/p>\n<p class=\"import-Normal\"><sup>2<\/sup> See Dunn Et al. 2016 and Rose Et al. 2018.<\/p>\n<p class=\"import-Normal\"><sup>3<\/sup> See Kirk and Williams 2011 and Seiffert Et al. 2009.<\/p>\n<p class=\"import-Normal\"><sup>4<\/sup> See Gregory 1920.<\/p>\n<p class=\"import-Normal\"><sup>5<\/sup> See Beard and MacPhee 1994 and Strait 2001.<\/p>\n<p class=\"import-Normal\"><sup>6<\/sup> See Schmid 1979.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\">\n<p class=\"import-Normal\">\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<p class=\"import-Normal\">Adapoids and omomyoids barely survived the Eocene-Oligocene extinctions, when colder temperatures, increased seasonality, and the retreat of rainforests to lower latitudes led to changes in mammalian biogeography. In North America, one genus (originally considered an omomyoid but recently reclassified as Adapoidea) persisted until the Miocene: <em>Ekgmowechashala<\/em> (Rose and Rensberger 1983). This taxon has highly unusual teeth and might have been a late immigrant to North America from Asia. In Asia, one family of adapoids, the Sivaladapidae, retained considerable diversity as late as the Late Miocene.<\/p>\n<h4 class=\"import-Normal\"><em>Adapoid Diversity<\/em><\/h4>\n<p class=\"import-Normal\">Adapoids were very diverse, particularly in the Eocene of North America and Europe. They can be divided into six families, with a few species of uncertain familial relationship. As a group, adapoids have some features in common, although much of what they share is plesiomorphic. Important features include the hallmarks of euprimates: postorbital bar, flattened nails, grasping extremities, and a petrosal bulla (Figures 9.9 and 9.10). In addition, some adapoids retain the ancestral dental formula of 2.1.4.3; that is, in each quadrant of the mouth, there are two incisors, one canine, four premolars, and three molars. In general, the incisors are small compared to the molars, but the canines are relatively large, with sexual dimorphism in some species. Cutting crests on the molars are well developed in some species, and the two halves of the mandible were fused at the midline in some species. Some adapoids were quite small (<em>Anchomomys <\/em>at a little over 100 g), and some were quite large (<em>Magnadapis<\/em> at 10 kg; 22 lbs.). Furthermore, the spaces and attachment features for the chewing muscles were truly enormous in some species, suggesting that these muscles were very large and powerful. Taken together, this suggests an overall adaptive profile of diurnal herbivory. The canine sexual dimorphism in some species suggests a possible mating pattern of polygyny, as males in polygynous primate species often compete with each other for mates and have especially large canine teeth.<\/p>\n<figure style=\"width: 548px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image18-1.jpg\" alt=\"Three partial animal crania.\" width=\"548\" height=\"350\" \/><figcaption class=\"wp-caption-text\">Figure 9.9: Representative crania of Adapidae from Museum d\u2019Histoire Naturelle Victor Brun, a natural history museum in Montauban, France. The white scale bar is 1 cm long. Credit: <a class=\"rId43\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Representative crania of adapids (European adapoids, (Figure 8.7)<\/a> from the <a class=\"rId44\" href=\"https:\/\/www.museum.montauban.com\/\">Museum d\u2019Histoire Naturelle Victor Brun in Montauban, France<\/a> original to <a class=\"rId45\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology <\/a>by Jonathan M. G. Perry is under a <a class=\"rId46\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<figure style=\"width: 547px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image19-2.jpg\" alt=\"Side views of small rodentlike skeleton with long tail.\" width=\"547\" height=\"525\" \/><figcaption class=\"wp-caption-text\">Figure 9.10: Darwinius masillae, a member of the Caenopithecidae. The slab on the left is Plate A and the slab on the right is Plate B. The parts of the skeleton in B that are outside of the dashed lines were fabricated. Credit: <a class=\"rId48\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Darwinius%20masillae%20holotype%20slabs.jpg\">Darwinius masillae holotype slabs<\/a> by Jens L. Franzen, Philip D. Gingerich, J\u00f6rg Habersetzer1, J\u00f8rn H. Hurum, Wighart von Koenigswald, B. Holly Smith is under a <a class=\"rId49\" href=\"https:\/\/creativecommons.org\/licenses\/by\/2.5\/legalcode\">CC BY 2.5 License<\/a>. Originally from Franzen Et al. 2009.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Omomyoid Diversity<\/em><\/h4>\n<p class=\"import-Normal\">Like adapoids, omomyoids appeared suddenly at the start of the Eocene and then became very diverse with most species dying out before the Oligocene. Omomyoids are known from thousands of jaws with teeth, relatively complete skulls for about a half-dozen species, and very little postcranial material. Omomyoids were relatively small primates, with the largest being less than three kilograms (approximately 7 lbs.; <em>Macrotarsius montanus<\/em>). All known crania possess a postorbital bar, which in some has been described as \u201cincipient closure.\u201d Some\u2014but not all\u2014known crania have an elongated bony ear tube extending lateral to the location of the eardrum, a feature seen in living tarsiers and <strong>catarrhines<\/strong>. The anterior teeth tend to be large, with canines that are usually not much larger than the incisors. Often it is difficult to distinguish closely related species using molar morphology, but the premolars tend to be distinct from one species to another. The postcranial skeleton of most omomyoids shows hallmarks of leaping behaviour reminiscent of that of tarsiers. In North America, omomyoids became very diverse and abundant. In fact, omomyoids from Wyoming are sufficiently abundant and from such stratigraphically controlled conditions that they have served as strong evidence for the gradual evolution of anatomical traits over time (Rose and Bown 1984).<\/p>\n<p class=\"import-Normal\"><em>Teilhardina <\/em>(Figure 9.11; see Figure 9.2) is one of the earliest and arguably the most plesiomorphic of omomyoids. <em>Teilhardina<\/em> has several species, most of which are from North America, with one from Europe (<em>T. belgica<\/em>) and one from Asia (<em>T. asiatica<\/em>). The species of this genus are anatomically similar and the deposits from which they are derived are roughly contemporaneous. Thus, this small primate likely dispersed across the northern continents very rapidly (Smith Et al. 2006).<\/p>\n<figure style=\"width: 545px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image3-1.jpg\" alt=\"World map with primates jumping across forested areas.\" width=\"545\" height=\"289\" \/><figcaption class=\"wp-caption-text\">Figure 9.11: A map of the world during the early Eocene showing one hypothesis for the direction of dispersal of the omomyoid Teilhardina. The map depicts primates hopping from continent to continent (East to West) via the forest corridors at high latitudes. Credit: <a href=\"https:\/\/www.pnas.org\/content\/103\/30\/11223\">Paleogeographic map showing hypothetical migration routes of Teilhardina (Figure 1)<\/a> by Thierry Smith, Kenneth D. Rose, and Philip D. Gingerich. 2006. <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">Proceedings of the National Academy of Sciences of the United States of America <\/a>103 (30): 11223\u201311227. Copyright (2006) National Academy of Sciences. Image <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">is used for non-commercial and educational purposes as outlined by PNAS.<\/a><\/figcaption><\/figure>\n<h2 class=\"import-Normal\">The Emergence of Modern Primate Groups<\/h2>\n<h3 class=\"import-Normal\"><strong>Origins of Crown Strepsirrhines<\/strong><\/h3>\n<p class=\"import-Normal\">Until the turn of this century, very little was known about the origins of the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1698\">crown<\/a><\/strong> (living) strepsirrhines. The Quaternary record of Madagascar contains many amazing forms of lemurs, including giant sloth-like lemurs, lemurs with perhaps monkey-like habits, lemurs with koala-like habits, and even a giant aye-aye (Godfrey and Jungers 2002). However, in Madagascar, early Tertiary continental sediments are lacking, and there is no record of lemur fossils before the Pleistocene.<\/p>\n<p class=\"import-Normal\">The fossil record of galagos is slightly more informative. Namely, there are Miocene African fossils that are very likely progenitors of lorisids (Simpson 1967). However, these are much like modern galagos and do not reveal anything about the relationship between crown strepsirrhines and Eocene fossil primates (but see below regarding <em>Propotto<\/em>). A similar situation exists for lorises in Asia: there are Miocene representatives, but these are substantially like modern lorises. The discovery of the first definite <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1699\"><strong>toothcomb<\/strong><\/a> canine (a hallmark of stresirrhines) in 2003 provided the \u201csmoking gun\u201d for the origin of crown strepsirrhines (Seiffert Et al. 2003). Recently, several other African primates have been recognized as having strepsirrhine affinities (Marivaux Et al. 2013; Seiffert 2012). The enigmatic Fayum primate <em>Plesiopithecus<\/em> is known from a skull that has been compared to aye-ayes and to lorises (Godinot 2006; Simons and Rasmussen 1994a).<\/p>\n<p class=\"import-Normal\">The now-recognized diversity of stem strepsirrhines from the Eocene and Oligocene of Afro-Arabia is strong evidence to suggest that strepsirrhines originated in that geographic area. This implies that lorises dispersed to Asia subsequent to an African origin. It is unknown what the first strepsirrhines in Madagascar were like. However, it seems likely that the lemuriform-lorisiform split occurred in continental Africa, followed by dispersal of lemuriform stock to Madagascar. Recent evidence suggests that <em>Propotto<\/em>, a Miocene primate from Kenya originally described as a potto antecedent, actually forms a clade with <em>Plesiopithecus <\/em>and the aye-aye; this might suggest that strepsirrhines dispersed to Madagascar from continental Africa more than once (Gunnell Et al. 2018).<\/p>\n<h3 class=\"import-Normal\"><strong>The Fossil Record of Tarsiers<\/strong><\/h3>\n<p class=\"import-Normal\">Tarsiers are so unusual that they fuel major debates about primate taxonomy. Tarsiers today are moderately diverse but geographically limited and not very different in their ecological habits\u2014especially considering that the split between them and their nearest living relative probably occurred over 50 million years ago. If omomyoids are excluded, then the fossil record of tarsiers is very limited. Two fossil species from the Miocene of Thailand have been placed in the genus <em>Tarsius<\/em>, as has an Eocene fossil from China (Beard Et al. 1994). These, and <em>Xanthorhysis<\/em> from the Eocene of China, are all very tarsier-like. In fact, it is striking that <em>Tarsius eocaenus<\/em> from China was already so tarsier-like as early as the Eocene. This suggests that tarsiers achieved their current morphology very early in their evolution and have remained more or less the same while other primates changed dramatically. Two additional genera, <em>Afrotarsius<\/em> from the Oligocene of Egypt and Libya and <em>Afrasia<\/em> from the Eocene of Myanmar, have also been implicated in tarsier origins, though the relationship between them and tarsiers is unclear (Chaimanee Et al. 2012). More recently, a partial skeleton of a small Eocene primate from China, <em>Archicebus achilles<\/em> (dated to approximately 55.8 million to 54.8 million years ago), was described as the most basal tarsiiform (Ni Et al. 2013). This primate is reconstructed as a diurnal insectivore and an arboreal quadruped that did some leaping\u2014but not to the specialized degree seen in living tarsiers. The anatomy of the eye in living tarsiers suggests that their lineage passed through a diurnal stage, so <em>Archicebus<\/em> (and diurnal omomyoids) might represent such a stage.<\/p>\n<h3 class=\"import-Normal\"><strong>Climate Change and the Paleogeography of Modern Primate Origins<\/strong><\/h3>\n<p class=\"import-Normal\">Changing global climate has had profound effects on primate dispersal patterns and ecological habits over evolutionary time. Primates today are strongly tied to patches of trees and particular plant parts such as fruits, seeds, and immature leaves. It is no surprise, then, that the distribution of primates mirrors the distribution of forests. Today, primates are most diverse in the tropics, especially in tropical rainforests. Global temperature trends across the Tertiary have affected primate ranges. Following the Cretaceous-Tertiary extinction event, cooler temperatures and greater seasonality characterized the Paleocene. In the Eocene, temperatures (and probably rainfall) increased globally and rainforests likely extended to very high latitudes. During this time, euprimates became diverse. With cooling and increased aridity at the end of the Eocene, many primate extinctions occurred in the northern continents and the surviving primates were confined to lower latitudes in South America, Afro-Arabia, Asia, and southern Europe. Among these survivors are the progenitors of the living groups of primates: lemurs and lorises, tarsiers, <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1700\"><strong>platyrrhines<\/strong><\/a> (monkeys of the Americas), and catarrhines (monkeys and apes of Africa and Asia) (Figure 9.12).<\/p>\n<figure id=\"attachment_277-9\" aria-describedby=\"caption-attachment-277-9\" style=\"width: 539px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-264\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image2-5-e1691791570984.png\" alt=\"Map of world with gray continents.\" width=\"539\" height=\"306\" \/><figcaption id=\"caption-attachment-277-9\" class=\"wp-caption-text\">Figure 9.12: Map of key localities of early anthropoids on land that becomes Africa and southern Asia. Credit: <a class=\"rId56\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Oligocene Map with Key Early Anthropoid Localities (Figure 8.10)<\/a> original to <a class=\"rId57\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Elyssa Ebding at <a class=\"rId58\" href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\">GeoPlace, California State University, Chico<\/a> is under a <a class=\"rId59\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Localities based on Fleagle 2013, 265.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Competing Hypotheses for the Origin of Anthropoids<\/strong><\/h3>\n<p class=\"import-Normal\">There is considerable debate among paleoanthropologists as to the geographic origins of anthropoids. In addition, there is debate regarding the source group for anthropoids. Three different hypotheses have been articulated in the literature. These are the adapoid origin hypothesis, the omomyoid origin hypothesis, and the tarsier origin hypothesis (Figure 9.13).<\/p>\n<figure style=\"width: 419px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image24-1-1.jpg\" alt=\"Diagrams show three relationships among primate groups.\" width=\"419\" height=\"742\" \/><figcaption class=\"wp-caption-text\">Figure 9.13: Competing models of anthropoid origins. Branch lengths are not to scale. The omomyoid origin model and tarsier origin model do not make specific reference to the evolutionary position of strepsirrhines; however, they were included here for completeness. <a href=\"https:\/\/docs.google.com\/document\/d\/1VUDKMBJYS_jNONjLxT04jQN0_z9Ua50BRN6auGSHUuU\/edit\">A full text description of this image is available<\/a>. Credit: <a class=\"rId61\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Competing Trees for Anthropoid Origins (Figure 8.11)<\/a> original to <a class=\"rId62\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Jonathan M. G. Perry is under a <a class=\"rId63\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Adapoid Origin Hypothesis<\/em><\/h4>\n<p class=\"import-Normal\">Resemblances between some adapoids and some extant anthropoids include fusion of the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1702\"><strong>mandibular symphysis<\/strong><\/a>, overall robusticity of the chewing system, overall large body size, features that signal a diurnal lifestyle (like relatively small eye sockets), and ankle bone morphology. Another feature in common is canine sexual dimorphism, which is present in some species of adapoids (probably) and in several species of anthropoids.<\/p>\n<p class=\"import-Normal\">These features led some paleoanthropologists in the last half of the 20th century to suggest that anthropoids came from adapoid stock (Gingerich 1980; Simons and Rasmussen 1994b). One of the earliest supporters of the link between adapoids and anthropoids was Hans Georg Stehlin, who described much of the best material of adapoids and compared these Eocene primates to South American monkeys (Stehlin 1912). In more recent times, the adapoid origin hypothesis was reinforced by resemblances between these European adapoids (especially <em>Adapis <\/em>and <em>Leptadapis<\/em>) and some early anthropoids from the Fayum Basin (e.g., <em>Aegyptopithecus<\/em>, see below; Figure 9.14).<\/p>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 473.25pt\">\n<caption>Figure 9.14: Families of early anthropoids with example genera and traits: a table. Credit: Early anthropoids table original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Jonathan M. G. Perry and Stephanie L. Canington is under a <a class=\"rId64\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Content derived from Fleagle 2013.<\/caption>\n<thead>\n<tr style=\"height: 25pt\">\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Family<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Genera<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Morphology<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Location<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Age<\/strong><sup><strong>1<\/strong><\/sup><\/p>\n<p>&nbsp;<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table3-R\" style=\"height: 18pt\">\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Propliopithecidae<sup>2<\/sup><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Aegyptopithecus<\/em><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Large size. Cranial sexual dimorphism, large canines. Robust jaws and rounded molars. Partially ossified ear tube (in some). Robust skeleton; quadruped.<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Africa<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Late Eocene to Early Oligocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 16pt\">\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Parapithecidae<sup>3<\/sup><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Apidium<\/em><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Medium size. Retention of three premolars per quadrant. Rounded molars and premolars with large central cusps. Adaptations for leaping in the lower limb.<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Africa<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Late Eocene to Late Oligocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 16pt\">\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Proteopithecidae<sup>4<\/sup><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Proteopithecus<\/em><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Small size. Retention of three premolars per quadrant. Arboreal quadrupeds that ate fruit.<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Africa<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Late Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 16pt\">\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Oligopithecidae<sup>5<\/sup><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Catopithecus<\/em><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Small size. Skull has postorbital septum and unfused mandible. Deep jaws. Diet of fruits. Generalized quadruped.<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Africa<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Late Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 16pt\">\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Eosimiidae<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Eosimias<\/em><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Deep jaw with vertical unfused symphysis. Pointed incisors and canines. Crowded premolars.<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Asia<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Middle Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 16pt\">\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Amphipithecidae<sup>6<\/sup><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Pondaungia<\/em><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Deep jaws. Molars generally rounded with wide basins.<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Asia<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Middle Eocene to Early Oligocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 1pt\">\n<td class=\"Table3-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\" colspan=\"4\">\n<p class=\"import-Normal\"><sup>1<\/sup> Derived from Fleagle 2013.<\/p>\n<p class=\"import-Normal\"><sup>2<\/sup> See Gebo and Simons 1987 and Simons Et al. 2007.<\/p>\n<p class=\"import-Normal\"><sup>3<\/sup> See Feagle and Simons 1995 and Simons 2001.<\/p>\n<p class=\"import-Normal\"><sup>4<\/sup> See Simons and Seiffert 1999.<\/p>\n<p class=\"import-Normal\"><sup>5<\/sup> See Simons and Rasmussen 1996.<\/p>\n<p class=\"import-Normal\"><sup>6<\/sup> See Kay Et al. 2004.<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\">\n<p class=\"import-Normal\">\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<p class=\"import-Normal\">Unfortunately for the adapoid hypothesis, most of the shared features listed above probably emerged independently in the two groups as adaptations to a diet of hard and\/or tough foods. For example, fusion of the mandibular symphysis likely evolved as a means to strengthen the jaw against forces that would pull the two halves away from each other, in the context of active chewing muscles on both sides of the head generating great bite forces. This context would also favour the development of robust jaws, large chewing muscles, shorter faces, and some other features shared by some adapoids and some anthropoids.<\/p>\n<p class=\"import-Normal\">As older and more plesiomorphic anthropoids were found in the Fayum Basin, it became clear that the earliest anthropoids from Africa did not possess these features of jaw robusticity (Seiffert Et al. 2009). Furthermore, many adapoids never evolved these features. Fusion of the mandibular symphysis in adapoids is actually quite different from that in anthropoids and probably occurred during juvenile development in the former (Beecher 1983; Ravosa 1996). Eventually, the adapoid origin hypothesis fell out of favour among most paleoanthropologists, although the description of <em>Darwinius<\/em> is a recent revival of that idea (Franzen Et al. 2009; but see Seiffert Et al. 2009, Williams Et al. 2010b).<\/p>\n<h4 class=\"import-Normal\"><em>Omomyoid Origin Hypothesis<\/em><\/h4>\n<p class=\"import-Normal\">Similarities in cranial and hindlimb morphology between some omomyoids and extant tarsiers have led to the suggestion that tarsiers arose from some kind of omomyoid. In particular, <em>Necrolemur<\/em> has many features in common with tarsiers, as does the North American <em>Shoshonius<\/em>, which is known from a few beautifully preserved (although distorted) crania. Tarsiers and <em>Shoshonius <\/em>share exclusively some features of the base of the cranium; however, <em>Shoshonius<\/em> does not have any sign of postorbital closure, and it lacks the bony ear tube of tarsiers. Nevertheless, some of the resemblances between some omomyoids and tarsiers suggest that tarsiers might have originated from within the Omomyoidea (Beard 2002; Beard and MacPhee 1994). In this scenario, although living tarsiers and living anthropoids might be sister taxa, they might have evolved from different omomyoids, possibly separated from each other by more than 50 million years of evolution, or from anthropoids evolved from some non-omomyoid fossil group. The arguments against the omomyoid origin hypothesis are essentially the arguments <em>for<\/em> the tarsier origin hypothesis (see below). Namely, tarsiers and anthropoids share many features (especially of the soft tissues) that must have been retained for many millions of years or must have evolved convergently in the two groups. Furthermore, a key hard-tissue feature shared between the two extant groups, the postorbital septum, was not present in any omomyoid. Therefore, that feature must have arisen convergently in the two extant groups or must have been lost in omomyoids. Neither scenario is very appealing, although recent arguments for <strong>convergent evolution<\/strong> of the postorbital septum in tarsiers and anthropoids have arisen from embryology and histology of the structure (DeLeon Et al. 2016).<\/p>\n<h4 class=\"import-Normal\"><em>Tarsier Origin Hypothesis<\/em><\/h4>\n<p class=\"import-Normal\">Several paleoanthropologists have suggested that there is a relationship between tarsiers and anthropoids to the exclusion of omomyoids and adapoids (e.g., Cartmill and Kay 1978; Ross 2000; Williams and Kay 1995). Tarsiers and anthropoids today share several traits, including many soft-tissue features related to the olfactory system (e.g., the loss of a hairless external nose and loss of the median cleft running from the nose to the mouth, as possessed by strepsirrhines), and aspects of the visual system (e.g., the loss of a reflective layer at the back of the eye, similarities in carotid circulation to the brain, and mode of placentation). Unfortunately, none of these can be assessed directly in fossils. Some bony similarities between tarsiers and anthropoids include an extra air-filled chamber below the middle ear cavity, reduced bones within the nasal cavity, and substantial postorbital closure; these can be assessed in fossils, but the distribution of these traits in omomyoids does not yield clear answers. Furthermore, several similarities between tarsiers and anthropoids are probably due to similarities in sensory systems, which might have evolved in parallel for ecological reasons. Although early attempts to resolve the crown primates with molecular data were sometimes equivocal or in disagreement with one another, more recent analyses (including those of short interspersed elements) suggest that tarsiers and anthropoids are sister groups to the exclusion of lemurs and lorises (Williams Et al. 2010a). However, this does not address omomyoids, all of which are far too ancient for DNA extraction.<\/p>\n<p class=\"import-Normal\">The above three hypotheses are not the only possibilities for anthropoid origins. It may be that anthropoids are neither the closest sister group of tarsiers, nor evolved from adapoids or omomyoids. In recent years, two new groups of Eocene Asian primates have been implicated in the origin of anthropoids: the eosimiids and the amphipithecids. It is possible that one or the other of these two groups gave rise to anthropoids. Regardless of the true configuration of the tree for crown primates, the three major extant groups probably diverged from each other quite long ago (Seiffert Et al. 2004).<\/p>\n<h3 class=\"import-Normal\"><strong>Early Anthropoid Fossils in Africa<\/strong><\/h3>\n<figure style=\"width: 526px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image7-2.jpg\" alt=\"People digging in a sandy desert.\" width=\"526\" height=\"352\" \/><figcaption class=\"wp-caption-text\">Figure 9.15: Egyptian workers sweeping Quarry I in the Fayum Basin (2004). This technique, called wind harvesting, removes the desert crust and permits wind to blow out fine sediment and reveal fossils. Credit: <a class=\"rId66\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Egyptian workers sweeping Quarry I in the Fayum Basin (2004, Figure 8.12)<\/a> by Jonathan M. G. Perry is under a <a class=\"rId67\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<figure style=\"width: 280px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image14-2.jpg\" alt=\"A person using a tool to expose bone in sand.\" width=\"280\" height=\"423\" \/><figcaption class=\"wp-caption-text\">Figure 9.16: Elwyn Laverne Simons excavating Aegyptopithecus in the Fayum Basin. Credit: <a class=\"rId69\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Elwyn Laverne Simons in the Fayum Basin (Figure 8.13)<\/a> used by permission of the <a class=\"rId70\" href=\"https:\/\/lemur.duke.edu\/\">Duke Lemur Center,<\/a> Division of Fossil Primates, is under a <a class=\"rId71\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">The classic localities yielding the greatest wealth of early anthropoid fossils are those from the Fayum Basin in Egypt (Simons 2008; Figure 9.15). The Fayum is a veritable oasis of fossil primates in an otherwise spotty early Tertiary African record. Since the 1960s, teams led by E. L. Simons have discovered several new species of early anthropoids, some of which are known from many parts of the skeleton and several individuals (Figure 9.16).<\/p>\n<p class=\"import-Normal\">The Fayum Jebel Qatrani Formation and Birket Qarun Formation between them have yielded a remarkable array of terrestrial, arboreal, and aquatic mammals. These include ungulates, bats, sea cows, elephants, hyraces, rodents, whales, and primates. Also, many other vertebrates, like water birds, were present. The area at the time of deposition (Late Eocene through Early Oligocene) was probably very wet, with slow-moving rivers, standing water, swampy conditions, and lots of trees (see Bown and Kraus 1988). In short, it was an excellent place for primates.<\/p>\n<h4 class=\"import-Normal\"><em>General Morphology of Anthropoids<\/em><\/h4>\n<p class=\"import-Normal\">The anthropoids known from the Fayum (and their close relatives from elsewhere in East Africa and Afro-Arabia) bear many of the anatomical hallmarks of extant anthropoids; however, there are plesiomorphic forms in several families that lack one or more anthropoid traits. All Fayum anthropoids known from skulls possess postorbital closure, most had fused mandibular symphyses, and most had ring-like <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1704\"><strong>ectotympanic<\/strong> <\/a> bones. Tooth formulae were generally either 2.1.3.3 or 2.1.2.3. Fayum anthropoids ranged in size from the very small <em>Qatrania<\/em> and <em>Biretia <\/em>(less than 500 g) to the much-larger <em>Aegyptopithecus<\/em> (approximately 7 kg; 15 lbs.). Fruit was probably the main component of the diet for most or all of the anthropoids, with some of them supplementing with leaves (Kay and Simons 1980; Kirk and Simons 2001; Teaford Et al. 1996). Most Fayum anthropoids were probably diurnal above-branch quadrupeds. Some of them (e.g., <em>Apidium<\/em>; see Figure 9.14) were probably very good leapers (Gebo and Simons 1987), but none show specializations for gibbon-style suspensory locomotion. Some of the Fayum anthropoids are known from hundreds of individuals, permitting the assessment of individual variation, sexual dimorphism, and in some cases growth and development. The description that follows provides greater detail for the two best known Fayum anthropoid families, the Propliopithecidae and the Parapithecidae; the additional families are summarized briefly.<\/p>\n<h4 class=\"import-Normal\"><em>Fayum Anthropoid Families<\/em><\/h4>\n<p class=\"import-Normal\">The Propliopithecidae (see Figure 9.14) include the largest anthropoids from the fauna, and they are known from several crania and some postcranial elements. They have been suggested to be stem catarrhines, although perhaps near the split between catarrhines and platyrrhines. The best known propliopithecid is <em>Aegyptopithecus<\/em>, known from many teeth, crania, and postcranial elements (Figure 9.17) .<\/p>\n<figure style=\"width: 431px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-2-1.jpg\" alt=\"Two animal skull side views.\" width=\"431\" height=\"281\" \/><figcaption class=\"wp-caption-text\">Figure 9.17: Female (left) and male (right) skull material for Aegyptopithecus zeuxis. The mandibles are not associated with the crania. Credit: <a href=\"https:\/\/www.pnas.org\/doi\/full\/10.1073\/pnas.0703129104#supplementary-materials\">Female and male cranium of A. zeuxi (03129Fig5, Supporting Information)<\/a> by Elwyn L. Simons, Erik R. Seiffert, Timothy M. Ryan, and Yousry Attia. 2007. <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">Proceedings of the National Academy of Sciences of the United States of America<\/a> 104 (21): 8731\u20138736. Copyright (2007) National Academy of Sciences. Image <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">is used for non-commercial and educational purposes as outlined by PNAS.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\">Parapithecidae are an extremely abundant and unusual family of anthropoids from the Fayum. The parapithecid <em>Apidium<\/em> is known from many jaws with teeth, crushed and distorted crania, and several skeletal elements. <em>Parapithecus<\/em> is known from cranial material including a beautiful, undistorted cranium. This genus shows extreme reduction of the incisors, including complete absence of the lower incisors in <em>P. grangeri <\/em>(Simons 2001). This trait is unique among primates. Parapithecids were once thought to be the ancestral stock of platyrrhines; however, their platyrrhine-like features are probably ancestral retentions, so the most conservative approach is to consider them stem anthropoids.<\/p>\n<p class=\"import-Normal\">The Proteopithecidae were small frugivores that probably mainly walked along horizontal branches on all fours. They are considered stem anthropoids. The best known genus, <em>Proteopithecus<\/em>, is represented by dentitions, crania, and post-cranial elements.<\/p>\n<p class=\"import-Normal\">The Oligopithecidae share a mixture of traits that makes them difficult to classify more specifically within anthropoids. The best known member, <em>Catopithecus<\/em>, is known from crania that demonstrate a postorbital septum and from mandibles that lack symphyseal fusion. They share the catarrhine tooth formula of 2.1.2.3 and have a canine honing complex that involves the anterior lower premolar. The postcranial elements known for the group suggest generalized arboreal quadrupedalism. The best known member, <em>Catopithecus<\/em>, is known from crania that demonstrate a postorbital septum and from mandibles that lack symphyseal fusion (Simons and Rasmussen 1996). The jaws are deep, with broad muscle attachment areas and crested teeth. <em>Catopithecus<\/em> was probably a little less than a kilogram in weight.<\/p>\n<p class=\"import-Normal\">Other genera of putative anthropoids from the Fayum include the very poorly known <em>Arsinoea<\/em>, the contentious <em>Afrotarsius<\/em>, and the enigmatic <em>Nosmips<\/em>. The last of these possesses traits of several major primate <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1705\">clades<\/a><\/strong> and defies classification (Seiffert Et al. 2010).<\/p>\n<h3 class=\"import-Normal\"><strong>Early Anthropoid Fossils in Asia<br style=\"clear: both\" \/><\/strong><\/h3>\n<p class=\"import-Normal\">For the last half of the 1900s, researchers believed that Africa was the unquestioned homeland of early anthropoids (see Fleagle and Kay 1994). However, two very different groups of primates from Asia soon began to change that. One was an entirely new discovery (Eosimiidae), and the other was a poorly known group discovered decades prior (Amphipithecidae). Soon, attention on anthropoid origins began to shift eastward (see Ross and Kay 2004; Simons 2004). If anthropoids arose in Asia instead of Africa, then this implies that the African early anthropoids either emigrated from Asia or evolved their anthropoid traits in parallel with living anthropoids.<\/p>\n<h4 class=\"import-Normal\"><em>Eosimiids<\/em><\/h4>\n<p class=\"import-Normal\">First described in the 1990s, the eosimiids are best represented by <em>Eosimias <\/em>(see Figure 9.14; Figure 9.18). This tiny \u201cdawn monkey\u201d is known from relatively complete jaws with teeth, a few small fragments of the face, and some postcranial elements (Beard Et al. 1994; Beard Et al. 1996; Gebo Et al. 2000). <em>Eosimias<\/em> (along with the other less-well-known genera in its family) bears some resemblance to tarsiers as well as anthropoids. Unfortunately, no good crania are known for this family, and the anatomy of, for example, the posterior orbital margin could be very revealing as to higher-level relationships.<\/p>\n<figure style=\"width: 550px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image16-1-1.jpg\" alt=\"Red-colored lower jaw of an animal.\" width=\"550\" height=\"232\" \/><figcaption class=\"wp-caption-text\">Figure 9.18: Cast of the right half of the mandible of Eosimias centennicus, type specimen. The white scale bar is 1 cm long. Credit: <a class=\"rId74\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Cast of the right half of the mandible of <\/a><a class=\"rId75\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\"><em>Eosimias centennicus <\/em><\/a><a class=\"rId76\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">(Figure 8.15),<\/a> type specimen, from K. D. Rose cast collection, photo by Jonathan M. G. Perry is under a <a class=\"rId77\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Amphipithecids<\/em><\/h4>\n<p class=\"import-Normal\">Amphipithecids are small- to medium-size primates (up to 10 kg; 22 lbs.). Most are from the Eocene Pondaung Formation in Myanmar (Early\u2013Middle Eocene), but one genus is known from Thailand. Some dental similarities with anthropoids were noted early on, such as deep jaws and wide basins that separate low molar cusps. The best known genera were <em>Pondaungia<\/em> and <em>Amphipithecus <\/em>(Ciochon and Gunnell 2002; see Figure 9.14). Another amphipithecid, <em>Siamopithecus<\/em> from Thailand, has very rounded molars and was probably a seed-eater (Figure 9.19). In addition to teeth and jaws, some cranial fragments, ankle material, and ends of post-cranial bones have been found for <em>Pondaungia<\/em>. There are important resemblances between these bones of <em>Pondaungia<\/em> and those of adapoids, suggesting adapoid affinities for the amphipithecidae. This would imply that the resemblances with anthropoids in the teeth are convergent, based on similarities in diet (see Ciochon and Gunnell 2002). Unfortunately, the association between post-cranial bones and teeth is not definite. With other primates in these faunas (including eosimiids), one cannot be certain that the post-cranial bones belong with the teeth. Some researchers suggest that some bones belong to a sivaladapid (or asiadapid) and others to an early anthropoid (Beard Et al. 2007; Marivaux Et al. 2003). Additional well-associated material of amphipithecids would help to clear up this uncertainty.<\/p>\n<figure style=\"width: 505px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image15-2.jpg\" alt=\"Four casts of jawbone fragments with teeth.\" width=\"505\" height=\"368\" \/><figcaption class=\"wp-caption-text\">Figure 9.19: Casts of representative amphipithecid material. A. Pondaungia cotteri right lower jaw fragment with m2 and m3. B. Siamopithecus eocaenus right upper jaw fragment with p4-m3. C. S. eocaenus right lower jaw fragment with partial m1, m2, and m3 in lateral view. D. Same as in C but occlusal view. White scale bars are 1 cm long. Credit: <a class=\"rId79\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Casts of representative amphipithecid material (Figure 8.16)l<\/a> from K. D. Rose cast collection, photo by Jonathan M. G. Perry is under a <a class=\"rId80\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Platyrrhine Dispersal to South America<\/strong><\/h3>\n<p class=\"import-Normal\">Today there is an impressive diversity of primates in South and Central America. These are considered to be part of a single clade, the Platyrrhini. Primates colonized South America sometime in the Eocene from an African source. In the first half of the 20th century, the source of platyrrhines was a matter of major debate among paleontologists, with some favouring a North American origin (e.g., Simpson 1940).<\/p>\n<p class=\"import-Normal\">Part of the reason for this debate is that South America was an island in the Eocene. Primates needed to cross open ocean to get there from either North America or Africa, although the distance from the former was shorter. Morphology yields clues to platyrrhine origins. The first known primates in South America have more in common morphologically with African primates than with North American ones. At the time, anthropoids were popping up in North Africa, whereas the only euprimates in North America were adapoids and omomyoids. Despite lacking a bony ear tube, early platyrrhines shared a great deal with other anthropoids, including full postorbital closure and fusion of the mandibular symphysis.<\/p>\n<p class=\"import-Normal\">The means by which a population of small North African primates managed to disperse across the Atlantic and survive to colonize South America remains a mystery. The most plausible scenario is one of rafting. That is, primates must have been trapped on vegetation that was blown out to sea by a storm. The vegetation then became a sort of life raft, which eventually landed ashore, dumping its passengers in South America. Rodents probably arrived in South America in the same way (Antoine Et al. 2012).<\/p>\n<p class=\"import-Normal\">Once ashore, platyrrhines must have crossed South America fairly rapidly because the earliest-known primates from that continent are from Peru (Bond Et al. 2015). Soon after that, platyrrhines were in Bolivia, namely <em>Branisella<\/em>. By the Miocene, platyrrhines were living in extreme southern Argentina and were exploiting a variety of feeding niches. The Early Miocene platyrrhines were all somewhat plesiomorphic in their morphology, but some features that likely arose by ecological convergence suggest (to some) relationships with extant platyrrhine families. This has led to a lively debate about the pattern of primate evolution in South America (Kay 2015; Kay &amp; Fleagle 2010; Rosenberger 2010). By the Middle Miocene, clear representatives of modern families were present in a diverse fauna from La Venta, Colombia (Wheeler 2010). The Plio-Pleistocene saw the emergence of giant platyrrhines as well as several taxa of platyrrhines living on Caribbean islands (Cooke Et al. 2016).<\/p>\n<p class=\"import-Normal\">The story of platyrrhines seems to be one of amazing sweepstakes dispersal, followed by rapid diversification and widespread geographic colonization of much of South America. After that, dramatic extinctions resulted in the current, much-smaller geographic distribution of platyrrhines. These extinctions were probably caused by changing climates, leading to the contraction of forests. Platyrrhines dispersed to the Caribbean and to Central America, with subsequent extinctions in those regions that might have been related to interactions with humans. Unlike anthropoids of Africa and Asia, platyrrhines do not seem to have evolved any primarily terrestrial forms and so have always been highly dependent on forests.<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Jonathan Perry and Primates of the Extreme South<\/h2>\n<p class=\"import-Normal\">Many primates are very vulnerable to ecological disturbance because they are heavily dependent on fruit to eat and trees to live in. This is one reason why so many primates are endangered today and why many of them went extinct due to climatic and vegetational changes in the past. I (Jonathan Perry) have conducted paleontological research focusing on primates that lived on the edge of their geographic distribution. This research has taken me to extreme environments in the Americas: southern Patagonia, the Canadian prairies, western Wyoming, and the badlands of eastern Oregon.<\/p>\n<p class=\"import-Normal\">Santa Cruz Province in Argentina is as far south as primates have ever lived. The Santa Cruz fauna of the Miocene has yielded a moderate diversity of platyrrhines, each with slightly different dietary adaptations. These include <em>Homunculus<\/em>, first described by Florentino Ameghino in 1891 (Figure 9.20). Recent fieldwork by my colleagues and I in Argentina has revealed several skulls of <em>Homunculus <\/em>as well as many parts of the skeleton (Kay Et al. 2012). The emerging profile of this extinct primate is one of a dedicated arboreal quadruped that fed on fruits and leaves. Many of the foods eaten by <em>Homunculus<\/em> must have been very tough and were probably covered and impregnated with grit; we suspect this because the cheek teeth are very worn down, even in young individuals, and because the molar tooth roots were very large, presumably to resist strong bite forces (Perry Et al. 2010, 2014).<\/p>\n<figure style=\"width: 497px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image9-2.jpg\" alt=\"An animal skull, a partial skull, and a fossil jaw with teeth.\" width=\"497\" height=\"634\" \/><figcaption class=\"wp-caption-text\">Figure 9.20: Representative specimens of Homunculus patagonicus. A. Adult cranium in lateral view. B. Adult cranium surface reconstructed from microCT scans, with the teeth segmented out. C. Juvenile cranium. White scale bars are 1cm long. Credit: <a class=\"rId82\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Representative specimens of <\/a><a class=\"rId83\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\"><em>Homunculus patagonicus <\/em><\/a><a class=\"rId84\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">(Figure 8.17)<\/a> photo by Jonathan M. G. Perry is under a <a class=\"rId85\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">I began working in Argentina while a graduate student at Duke University. I participated as a field assistant in a team led by my Ph.D. advisor, Richard F. Kay, and Argentine colleagues Sergio F. Vizca\u00edno and M. Susana Bargo. Most of the localities examined belong to a suite of beach sites known since the 1800s and visited by many field parties from various museums in the early 1900s. Since 2003, our international team of paleontologists from the U.S. and Argentina has visited these localities every single year (Figure 9.21). Over time, new fossils and new students have led to new projects and new approaches, including the use of microcomputed tomography (microCT) to visualize and analyse internal structures of the skeleton.<\/p>\n<figure style=\"width: 491px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image23.jpg\" alt=\"Sandy rocky coastline. People digging on a grassy hillside.\" width=\"491\" height=\"561\" \/><figcaption class=\"wp-caption-text\">Figure 9.21: Field localities in Argentina and Canada. A. Ca\u00f1adon Palos locality, coastal Santa Cruz Province, Argentina. B. Swift Current Creek locality, southwest Saskatchewan, Canada. Credits: A. <a class=\"rId87\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Ca\u00f1adon Palos Field Locality in Argentina<\/a> by Jonathan M. G. Perry is under a <a class=\"rId88\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. B. <a class=\"rId89\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Swift Current Creek locality, Saskatchewan, Canada<\/a> by Jonathan M. G. Perry is under a <a class=\"rId90\" 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\">Planet of Apes<\/h2>\n<h3 class=\"import-Normal\"><strong>Geologic Activity and Climate Change in the Miocene<\/strong><\/h3>\n<p class=\"import-Normal\">The Miocene Epoch was a time of mammalian diversification and extinction, global climate change, and ecological turnover. In the Miocene, there was an initial warming trend across the globe with the expansion of subtropical forests, followed by widespread cooling and drying with the retreat of tropical forests and replacement with more open woodlands and eventually grasslands. It was also a time of major geologic activity. On one side of the globe, South America experienced the rise of the Andes Mountains. On the other side, the Indian subcontinent collided with mainland Asia, resulting in the rise of the Himalayan Mountains. In Africa, volcanic activity promoted the development of the East African Rift System. Critical to the story of ape evolution was the exposure of an intercontinental landbridge between East Africa and Eurasia, permitting a true planet of apes (Figure 9.22).<\/p>\n<figure id=\"attachment_277-10\" aria-describedby=\"caption-attachment-277-10\" style=\"width: 580px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-273\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image21-3-e1691792198797.png\" alt=\"Map of world with gray continents.\" width=\"580\" height=\"335\" \/><figcaption id=\"caption-attachment-277-10\" class=\"wp-caption-text\">Figure 9.22: Map of the world in the Miocene, highlighting fossil ape localities across Africa, southern Europe, and southern Asia. Credit: <a class=\"rId92\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Miocene Map with Fossil Ape Localities (Figure 8.19)<\/a> original to <a class=\"rId93\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Elyssa Ebding at <a class=\"rId94\" href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\">GeoPlace, California State University, Chico<\/a> is under a <a class=\"rId95\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Localities based on Fleagle 2013, 311.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Geographic Distribution: Africa, Asia, Europe<\/strong><\/h3>\n<p class=\"import-Normal\">The world of the Miocene had tremendous ape diversity compared to today. The earliest records of fossil apes are from Early Miocene deposits in Africa. However, something dramatic happened around 16 million years ago. With the closure of the ancient Tethys Sea, the subsequent exposure of the <em>Gomphotherium<\/em> Landbridge, and a period of global warming, the Middle\u2013Late Miocene saw waves of emigration of mammals (including primates) out of Africa and into Eurasia, with evidence of later African re-entry for some (Harrison 2010). Some of the mammals that dispersed from Africa to Eurasia and back were apes. Though most of these early apes left no modern descendants, some of them gave rise to the ancestors of modern apes\u2014including <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_800\">hominins<\/a><\/strong> (Figure 9.23).<\/p>\n<figure style=\"width: 560px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image20-1.jpg\" alt=\"Miocene apes set against a geologic time scale.\" width=\"560\" height=\"796\" \/><figcaption class=\"wp-caption-text\">Figure 9.23: Representative Miocene apes set against a geologic time scale. Credit: <a href=\"https:\/\/www.pnas.org\/content\/108\/14\/5554\">Range chart for Miocene hominoids of Western Eurasia (Figure 3)<\/a> by Isaac Casanovas-Vilar, David M. Alba, Miguel Garc\u00e9s, Josep M. Robles, and Salvador Moy\u00e0-Sol\u00e0. 2011. <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">Proceedings of the National Academy of Sciences of the United States of America<\/a> 108 (14): 5554-5559. Copyright (2011) National Academy of Sciences. Image <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">is used for non-commercial and educational purposes as outlined by PNAS.<\/a><\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Where Are the Monkeys? Diversity in the Miocene<\/strong><\/h3>\n<p class=\"import-Normal\">Whereas the Oligocene deposits in the Fayum of Egypt have yielded the earliest-known catarrhine fossils, the Miocene demonstrates some diversification of Cercopithecoidea. However, compared to the numerous and diverse Miocene apes (see below), monkeys of the Miocene are very rare and restricted to a single extinct family, the Victoriapithecidae (Figure 9.24). This family contains the earliest definite cercopithecoids.<\/p>\n<figure style=\"width: 357px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image13-5.png\" alt=\"Front view of skull with pointed teeth.\" width=\"357\" height=\"322\" \/><figcaption class=\"wp-caption-text\">Figure 9.25: Skull of Victoriapithecus macinnesi. Credit: Victoriapithecus macinnesi skull photo taken at the Musee d'Histoire Naturelle, Paris by Ghedoghedo is under a CC BY-SA 3.0 License.<\/figcaption><\/figure>\n<p class=\"import-Normal\">These monkeys are found from northern and eastern Africa between 20 million and 12.5 million years ago (Miller Et al. 2009). The best known early African monkey is <em>Victoriapithecus <\/em>(Figure 9.25), a small-bodied (approximately 7 kg; 15 lbs.), small-brained monkey. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1708\">Bilophodonty<\/a><\/strong>, known to be a hallmark of molar teeth of modern cercopithecoid, was present to some extent in Victoriapithecids. <em>Victoriapithecus<\/em> has been reconstructed as being more frugivorous and perhaps spent more time on the ground (terrestrial locomotion) than in the trees (arboreal locomotion; Blue Et al. 2006). The two major groups of cercopithecoids today are cercopithecines and colobines. The earliest records demonstrating clear members of each of these two groups are at the end of the Miocene. Examples include the early colobine <em>Microcolobus<\/em> from Kenya and the early cercopithecine <em>Pliopapio<\/em> from Ethiopia.<\/p>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 473.25pt;height: 349px\">\n<caption>Figure 9.24: Some families of later anthropoids with example genera and traits: a table. Credit: Late anthropoids table original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Jonathan M. G. Perry and Stephanie L. Canington is under a <a class=\"rId100\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Content derived from Fleagle 2013.<\/caption>\n<thead>\n<tr style=\"height: 25pt\">\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 60px;width: 119.35px\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Family<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 60px;width: 103.417px\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Genera<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 60px;width: 191.65px\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Morphology<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 60px;width: 67.3667px\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Location<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 60px;width: 73.2167px\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Age<\/strong><sup><strong>1<\/strong><\/sup><\/p>\n<p>&nbsp;<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table4-R\" style=\"height: 18pt\">\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 77px;width: 119.35px\">\n<p class=\"import-Normal\">Victoriapithecidae<sup>2<\/sup><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 77px;width: 103.417px\">\n<p class=\"import-Normal\"><em>Victoriapithecus<\/em><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 77px;width: 191.65px\">\n<p class=\"import-Normal\">Long, sloping face. Round, narrowly spaced orbits. Deep cheek bones. Well-developed sagittal crest.<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 77px;width: 67.3667px\">\n<p class=\"import-Normal\">Africa<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 77px;width: 73.2167px\">\n<p class=\"import-Normal\">Early to Middle Miocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 16pt\">\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 61px;width: 119.35px\">\n<p class=\"import-Normal\">Proconsulidae<sup>3<\/sup><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 61px;width: 103.417px\">\n<p class=\"import-Normal\"><em>Proconsul<\/em><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 61px;width: 191.65px\">\n<p class=\"import-Normal\">Short face. Generalized dentition. Arboreal quadruped. Probably tailless.<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 61px;width: 67.3667px\">\n<p class=\"import-Normal\">Africa and Arabia<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 61px;width: 73.2167px\">\n<p class=\"import-Normal\">Early to Middle Miocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 16pt\">\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 46px;width: 119.35px\">\n<p class=\"import-Normal\">Pongidae<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 46px;width: 103.417px\">\n<p class=\"import-Normal\"><em>Gigantopithecus<\/em><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 46px;width: 191.65px\">\n<p class=\"import-Normal\">Largest primate ever. Deep jaws and low rounded molars.<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 46px;width: 67.3667px\">\n<p class=\"import-Normal\">Asia<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 46px;width: 73.2167px\">\n<p class=\"import-Normal\">Miocene to Present<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 1pt\">\n<td class=\"Table4-C\" style=\"border-color: #000000;border-style: solid none none;border-width: 0.5pt 0pt 0pt;padding: 0pt 5.4pt;height: 90px;width: 526.983px\" colspan=\"4\">\n<p class=\"import-Normal\"><sup>1<\/sup> Derived from Fleagle 2013.<\/p>\n<p class=\"import-Normal\"><sup>2<\/sup> See Benefit and McCrossin 1997 and Fleagle 2013.<\/p>\n<p class=\"import-Normal\"><sup>3<\/sup> See Begun 2007.<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"border-color: #000000;border-style: solid none none;border-width: 0.5pt 0pt 0pt;padding: 0pt 5.4pt;height: 90px;width: 73.2167px\">\n<p class=\"import-Normal\">\n<\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"height: 15px;width: 121.283px\"><\/td>\n<td style=\"height: 15px;width: 105.35px\"><\/td>\n<td style=\"height: 15px;width: 193.583px\"><\/td>\n<td style=\"height: 15px;width: 69.3px\"><\/td>\n<td style=\"height: 15px;width: 74.65px\"><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<h3><\/h3>\n<h3><\/h3>\n<h3><\/h3>\n<h3 class=\"import-Normal\"><strong>The Story of Us, the Apes<\/strong><\/h3>\n<h4 class=\"import-Normal\"><em>African Ape Diversity <\/em><\/h4>\n<p class=\"import-Normal\">The Early Miocene of Africa has yielded around 14 genera of early apes (Begun 2003). Many of these taxa have been reconstructed as frugivorous arboreal quadrupeds (Kay 1977). One of the best studied of these genera is the East African <em>Proconsul<\/em> (Family Proconsulidae; see Figure 9.24). Several species have been described, with body mass reconstructions ranging from 17 to 50 kg (approximately 37\u2013110 lbs.). A paleoenvironmental study reconstructed the habitat of <em>Proconsul <\/em>to be a dense, closed-canopy tropical forest (Michel Et al. 2014). No caudal vertebrae (tail bones) have been found in direct association with <em>Proconsul <\/em>post-crania, and the morphology of the sacrum is consistent with <em>Proconsul<\/em> lacking a tail (Russo 2016; Ward Et al. 1991).<\/p>\n<p class=\"import-Normal\">Overall, the African ape fossil record in the Late Miocene is sparse, with seven fossil localities dating between eleven and five million years ago (Pickford Et al. 2009). Nevertheless, most species of great apes live in Africa today. Where did the progenitors of modern African apes arise? Did they evolve in Africa or somewhere else? The paucity of apes in the Late Miocene of Africa stands in contrast to the situation in Eurasia. There, ape diversity was high. Furthermore, several Eurasian ape fossils show morphological affinities with modern hominoids (apes). Because of this, some paleoanthropologists suggest that the ancestors of modern African great apes recolonized Africa from Eurasia toward the end of the Miocene (Begun 2002). However, discoveries of Late Miocene hominoids like the Kenyan <em>Nakalipithecus<\/em> (9.9 million to 9.8 million years ago), the Ethiopian <em>Chororapithecus<\/em> (10.7 million to 10.1 million years ago), and the late-Middle Miocene Namibian <em>Otavipithecus<\/em> (13 million to 12 million years ago) fuel an alternative hypothesis\u2014namely that African hominoid diversity was maintained throughout the Miocene and that one of these taxa might, in fact, be the last common ancestor of extant African apes (Kunimatsu Et al. 2007; Mocke Et al. 2002). The previously underappreciated diversity of Late Miocene apes in Africa might be due to poor sampling of the fossil record in Africa.<\/p>\n<h4 class=\"import-Normal\"><em>Eurasian Ape Diversity<\/em><\/h4>\n<p class=\"import-Normal\">With the establishment of the <em>Gomphotherium<\/em> Landbridge (a result of the closure of the Eastern Mediterranean seaway; R\u00f6gl 1999), the Middle Miocene was an exciting time for hominoid radiations outside of Africa (see Figure 9.23). Eurasian hominoid species exploited their environments in many different ways in the Miocene. Food exploitation ranged from soft-fruit feeding in some taxa to hard-object feeding in others, in part owing to seasonal fluctuations and the necessary adoptions of fallback foods (DeMiguel Et al. 2014). For example, the molars of <em>Oreopithecus bambolii<\/em> (Family Hominidae) have relatively long lower-molar shearing crests, suggesting that this hominoid was very folivorous (Ungar and Kay 1995). Associated with variation in diet, there is great variation in the degree to which cranial features (e.g., zygomatic bone or supraorbital tori) are developed across the many taxa (Cameron 1997); however, Middle Miocene fossils tend to exhibit relatively thick molar enamel and relatively robust jaws (Andrews and Martin 1991).<\/p>\n<p class=\"import-Normal\">In Spain, the cranium with upper dentition, part of a mandible, and partial skeleton of <em>Pliobates <\/em>(Family Pliobatidae), a small-bodied ape (4\u20135 kg; 9\u201311 lbs.), was discovered in deposits dating to 11.6 million years ago (Alba Et al. 2015). It is believed to be a frugivore with a relative brain size that overlaps with modern cercopithecoids. The fossilized postcrania of <em>Pliobates<\/em> suggest that this ape might have had a unique style of locomotion, including the tendency to walk across the branches of trees with its palms facing downward and flexible wrists that permitted rotation of the forearm during climbing. However, the anatomy of the distal humerus differs from those of living apes in ways that suggest that <em>Pliobates<\/em> was less efficient at stabilizing its elbow while suspended (Benefit and McCrossin 2015). Two other recently described apes from Spain, <em>Pierolapithecus <\/em>and <em>Anoiapithecus<\/em>, are known from relatively complete skeletons. <em>Pierolapithecus<\/em> had a very projecting face and thick molar enamel as well as some skeletal features that suggest (albeit controversially) a less suspensory locomotor style than in extant apes (Moy\u00e0-Sol\u00e0 Et al. 2004). In contrast to <em>Pierolapithecus<\/em>, the slightly younger <em>Anoiapithecus<\/em> has a very flat face (Moy\u00e0-Sol\u00e0 Et al. 2009).<\/p>\n<p class=\"import-Normal\">Post-cranial evidence for suspensory or well-developed orthograde behaviours in apes does not appear until the Late Miocene of Europe. Primary evidence supporting these specialized locomotor modes includes the relatively short lumbar vertebrae of <em>Oreopithecus <\/em>(Figure 9.26) and <em>Dryopithecus<\/em> (Maclatchy 2004). Further, fossil material of the lower torso of <em>O. bambolii <\/em>(which dates to the <em>Pan<\/em>-hominin divergence) conveys a higher degree of flexion-extension abilities in the lumbar region (lower back) than what is possible in extant apes. Additionally, the hindlimb of <em>O. bambolii <\/em>is suggested to have supported powerful hip adduction during climbing (Hammond Et al. 2020). The Late Miocene saw the extinction of most of the Eurasian hominoids in an event referred to as the Vallesian Crisis (Agust\u00ed Et al. 2003). Among the latest surviving hominoid taxa in Eurasia were <em>Oreopithecus<\/em> and <em>Gigantopithecus<\/em>, the latter of which held out until the Pleistocene in Asia and was probably even sympatric with <em>Homo erectus<\/em> (Cachel 2015).<\/p>\n<figure style=\"width: 436px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image8-2-1.jpg\" alt=\"Posterior view of ancient ape skeleton.\" width=\"436\" height=\"775\" \/><figcaption class=\"wp-caption-text\">Figure 9.26: Skeleton of Oreopithecus bambolii. Credit: <a class=\"rId107\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Oreopithecus_bambolii_1.JPG\">Oreopithecus bambolii 1<\/a> by <a class=\"rId108\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Ghedoghedo\">Ghedoghedo<\/a> is under a <a class=\"rId109\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>The Origins of Extant Apes<\/strong><\/h3>\n<p class=\"import-Normal\">The fossil record of the extant apes is somewhat underwhelming: it ranges from being practically nonexistent for some taxa (e.g., chimpanzees) to being a little better for others (e.g., humans). There are many possible reasons for these differences in fossil abundance, and many are associated with the environmental conditions necessary for the fossilization of bones. One way to understand the evolution of extant apes that is not so dependent on the fossil record is via molecular evolutionary analyses. This can include counting up the differences in the genetic sequence between two closely related species to estimate the amount of time since these species shared a common ancestor. This is called a molecular clock, and it is often calibrated using fossils of known absolute age that stand in for the last common ancestor of a particular clade. Molecular clock estimates have placed the Hylobatidae and Hominidae split between 19.7 million and 24.1 million years ago, the African ape and Asian ape split between 15.7 million and 19.3 million years ago, and the split of Hylobatidae into its current genera between 6.4 million and 8 million years ago (Israfil Et al. 2011).<\/p>\n<h4 class=\"import-Normal\"><em>Small Ape Origins and Fossils<\/em><\/h4>\n<p class=\"import-Normal\">Unfortunately, the fossil record for the small (formerly \u201clesser\u201d) apes is meager, particularly in Miocene deposits. One possible early hylobatid is <em>Laccopithecus robustus<\/em>, a Late Miocene catarrhine from China (Harrison 2016). Although it does share some characteristics with modern gibbons and siamangs (including an overall small body size and a short face), <em>Laccopithecus<\/em> most likely represents a plesiomorphic stem catarrhine and is therefore distantly related to extant apes (Jablonski &amp; Chaplin 2009). A more likely candidate for the hylobatid stem is another Late Miocene taxon from China, <em>Yuanmoupithecus xiaoyuan<\/em>. Interpretation of its phylogenetic standing, however, is complicated by contradicting dental features\u2014some of them quite plesiomorphic\u2014which some believe best place <em>Yuanmoupithecus<\/em> as a stem hylobatid (Harrison 2016). Recently, a Middle Miocene Indian fossil ape, <em>Kapi ramnagarensis<\/em>, has extended the fossil record of small apes by approximately five million years. Its teeth are suggestive of a shift to a more frugivorous diet and it is likely a stem hylobatid (Gilbert Et al. 2020). The history of Hylobatidae becomes clearer in the Pleistocene, with fossils representing extant genera.<\/p>\n<h4 class=\"import-Normal\"><em>Great Ape Origins and Fossils<\/em><\/h4>\n<p class=\"import-Normal\">The most extensive fossil record of a modern great ape is that of our own genus, <em>Homo<\/em>. However, the evolutionary history of the Asian great ape, the orangutan (<em>Pongo<\/em>), is becoming clearer. Today, orangutans are found only on the islands of Borneo and Sumatra. However, Pleistocene-aged teeth, attributed to <em>Pongo<\/em>, have been found in Cambodia, China, Laos, Peninsular Malaysia, and Vietnam\u2014demonstrating the vastness of the orangutan\u2019s previous range (Ibrahim Et al. 2013; Wang Et al. 2014). <em>Sivapithecus <\/em>from the Miocene of India and Pakistan is represented by many specimens, including parts of the face. <em>Sivapithecus<\/em> is very similar to <em>Pongo<\/em>, especially in the face, and it probably is closely related to ancestral orangutans (Pilbeam 1982). Originally, jaws and teeth belonging to the former genus <em>Ramapithecus<\/em> were thought to be important in the origin of humans (Simons 1961), but now these are recognized as specimens of <em>Sivapithecus<\/em> (Kelley 2002). Postcranial bones of <em>Sivapithecus<\/em>, however, suggest a more generalized locomotor mode\u2014including terrestrial locomotion\u2014than seen in <em>Pongo <\/em>(Pilbeam Et al. 1990). Stable carbon and oxygen isotope data from dental enamel have reconstructed the paleoecological space of <em>Sivapithecus <\/em>(as well as the contemporaneous Late Miocene pongine <em>Khoratpithecus<\/em>) within the canopies of forested habitats (Habinger Et al. 2022).<\/p>\n<p class=\"import-Normal\">A probable close relative of <em>Sivapithecus <\/em>is the amazing <em>Gigantopithecus<\/em> (see Figure 9.24). Known only from teeth and jaws from China and India (e.g., Figure 9.27), this ape probably weighed as much as 270 kg (595 lbs.) and was likely the largest primate ever (Bocherens Et al. 2017). Because of unique features of its teeth (including molarized premolars and patterns of wear) and its massive size, it has been reconstructed as a bamboo specialist, somewhat like the modern panda. Small silica particles (phytoliths) from grasses have been found stuck to the molars of <em>Gigantopithecus<\/em> (Ciochon Et al. 1990). Recent studies evaluating the carbon isotope composition of the enamel sampled from <em>Gigantopithecus<\/em> teeth suggest that this ape exploited a wide range of vegetation, including fruits, leaves, roots, and bamboo (Bocherens Et al. 2017). Its face is reminiscent of that of modern orangutans and it might belong in the same family, Pongidae (Kelley 2002).<\/p>\n<figure style=\"width: 488px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image12.jpg\" alt=\"Superior view of mandible and teeth.\" width=\"488\" height=\"533\" \/><figcaption class=\"wp-caption-text\">Figure 9.27: Cast of the mandible of Gigantopithecus blacki. Credit: <a class=\"rId111\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Gigantopithecus%20blacki%20mandible%20010112.jpg\">Gigantopithecus blacki mandible 010112<\/a> by <a class=\"rId112\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Wilson44691\">Wilson44691<\/a> is under a <a class=\"rId113\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">In Africa, the first fossil to be confidently attributed to <em>Pan<\/em>, and known to be the earliest evidence of a chimpanzee, was described based on teeth found in Middle Pleistocene deposits in the Eastern Rift Valley of Kenya (McBrearty and Jablonski 2005). Paleoenvironmental reconstructions of this locality suggest that this early chimpanzee was living in close proximity to early <em>Homo<\/em> in a closed-canopy wooded habitat. Similarly, fossil teeth and mandibular remains attributed to two species of Middle-Late Miocene apes\u2014<em>Chororapithecus abyssinicus<\/em> (from Ethiopia; Suwa Et al. 2007) and <em>Nakalipithecus nakayamai<\/em> (from Kenya; Kunimatsu Et al. 2007)\u2014have been suggested as basal members of the gorilla clade.<\/p>\n<p class=\"import-Normal\">While the deposits of Eastern Africa have yielded a profound record of our fossil hominin ancestors, the continent\u2019s rainforests remain a \u201cpalaeontological desert\u201d (Rosas Et al. 2022). Clearly, more work is needed to fill in the large gaps in the fossil record of the nonhuman great apes. The twentieth century witnessed the discovery of many hominin fossils in East Africa, which have been critical for improving our understanding of human evolution. While twenty-first-century conservationists fight to prevent the extinction of the living great apes, perhaps efforts by twenty-first-century paleoanthropologists will yield the evolutionary story of these, our closest relatives.<\/p>\n<h2>Cultural Connections and Contemporary Implications<\/h2>\n<p>Even today, continued study of primate evolution and the gathering of new evidence are critical to our understanding of our biological history. The Piltdown Man hoax is a striking example of how, without modern carbon dating technology and continued study in the field, false conclusions and evidence can be accepted as truth. In 1912, Charles Dawson\u2014an amateur antiquarian and archeologist\u2014presented a combination of human skull fragments and an ape-like lower jaw as evidence of a previously unknown \u2018missing link\u2019 in the line of human ancestors (Groote et al., 2016). Immediately, members of the British archaeological establishment accepted the specimen because it seemed consistent with the belief that a larger\u2014human-like\u2014brain was among the earliest features to evolve in humans. This specimen remained largely accepted for decades until, in 1953, modern chemical analyses proved that the remains had been deliberately altered: the jaw had been stained, the teeth filed, and the skull fragments were subsequently carbon-dated to only approximately 600 years old (Australian Museum, n.d.).<\/p>\n<p>Among many such cases, the Piltdown Man hoax demonstrates why the continued study of primate evolution remains imperative to contemporary biological anthropology. As new discoveries, technologies, and analytical methods provide additional evidence, our understanding of human biological history can be continually evaluated and refined.<\/p>\n<p>Discussion questions<\/p>\n<ul>\n<li style=\"font-weight: 400\">Can you think of an example where a belief that was widely accepted as scientific truth became harmful, not only to science but also to the people within that society?<\/li>\n<li style=\"font-weight: 400\">Why is it important for scientists to remain open to revising their conclusions when new evidence emerges? How does the Piltdown Man hoax demonstrate the importance of this?<\/li>\n<\/ul>\n<div class=\"textbox shaded\">\n<h2>Summary<\/h2>\n<p>While there are large gaps in the fossil records linking primates to early hominins, evolutionary trends make it clear that humans are one branch of the broader primate family tree. In this chapter we go over the major development that characterize primate evolution: enhanced vision, grasping hands and feet, greater reliance on social behaviour, and increased brain complexity. It is these traits which distinguish primates from other mammals and furthermore, help define major subsections within the primate class, such as strepsirrhines, haplorhines, monkeys, apes, and ultimately hominins.<\/p>\n<p>Within this chapter, we also examine how anthropologists reconstruct these evolutionary relationships. Fossil evidence has provided key information about when and where different primates lived, while genetic data such as skeletal features allow researchers to understand how extinct species moved, ate, and interacted with their environments. Just as crucial are the influences of Earth\u2019s changing environments: continental drift, glacial cycles, and long-term climate shifts have repeatedly reshaped habitats, driving both extinctions and the emergence of new adaptive forms, including the emergence of our own human lineage.<\/p>\n<h2 class=\"import-Normal\">Review Questions<strong><br \/>\n<\/strong><\/h2>\n<ul>\n<li>Compare three major hypotheses about primate origins, making reference to each one\u2019s key ecological reason for primate uniqueness.<\/li>\n<li>Explain how changes in temperature, rainfall, and vegetation led to major changes in primate biogeography over the Early Tertiary.<\/li>\n<li>List some euprimate features that plesiadapiforms have and some that they lack.<\/li>\n<li>Contrast adapoids and omomyoids in terms of life habits.<\/li>\n<li>Describe one piece of evidence for each of the adapoid, omomyoid, and tarsier origin hypotheses for anthropoids.<\/li>\n<li>Discuss the biogeography of the origins of African great apes and orangutans using examples from the Miocene ape fossil record.<\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\">Key Terms<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\"><strong>Adapoidea<\/strong>: Order: Primates. One of the earliest groups of euprimates (true primates; earliest records from the early Eocene).<\/p>\n<p class=\"import-Normal\"><strong>A<\/strong><strong>daptive radiations<\/strong>: Rapid diversifications of single lineages into many species which may present unique morphological features in response to different ecological settings.<\/p>\n<p class=\"import-Normal\"><strong>Ancestral traits<\/strong>: Features that were inherited from a common ancestor and which remain (largely) unchanged.<\/p>\n<p class=\"import-Normal\"><strong>Anthropoids<\/strong>:Group containing monkeys and apes, including humans.<\/p>\n<p class=\"import-Normal\"><strong>Auditory bulla<\/strong>: The rounded bony floor of the middle ear cavity.<\/p>\n<p class=\"import-Normal\"><strong>Bilophodonty<\/strong>: Dental condition in which the cusps of molar teeth form ridges (or lophs) separated from each other by valleys (seen, e.g., in modern catarrhine monkeys).<\/p>\n<p class=\"import-Normal\"><strong>Catarrhines<\/strong>: Order: Primates; Suborder: Anthropoidea; Infraorder: Catarrhini. Group, with origins in Africa and Asia, that contains monkeys and apes, including humans.<\/p>\n<p class=\"import-Normal\"><strong>Clade<\/strong>:Group containing all of the descendants of a single ancestor. A portion of a phylogenetic tree represented as a bifurcation (node) in a lineage and all of the branches leading forward in time from that bifurcation.<\/p>\n<p class=\"import-Normal\"><strong>Convergent evolution<\/strong>: The independent evolution of a morphological feature in animals not closely related (e.g., wings in birds and bats).<\/p>\n<p class=\"import-Normal\"><strong>Crown<\/strong>: Smallest monophyletic group (clade) containing a specified set of extant taxa and all descendants of their last common ancestor.<\/p>\n<p class=\"import-Normal\"><strong>Diastema<\/strong>: Space between adjacent teeth.<\/p>\n<p class=\"import-Normal\"><strong>Diffuse coevolution<\/strong>: The ecological interaction between whole groups of species (e.g., primates) with whole groups of other species (e.g., fruiting trees).<\/p>\n<p class=\"import-Normal\"><strong>Ectotympanic<\/strong>: Bony ring or tube that holds the tympanic membrane (eardrum).<\/p>\n<p class=\"import-Normal\"><strong>Euprimates<\/strong>: Order: Primates. True primates or primates of modern aspect.<\/p>\n<p class=\"import-Normal\"><strong>Haplorhines<\/strong>: Group containing catarrhines, platyrrhines, and tarsiers.<\/p>\n<p class=\"import-Normal\"><strong>Hominins<\/strong>: Modern humans and any extinct relatives more closely related to us than to chimpanzees.<\/p>\n<p class=\"import-Normal\"><strong>Mandibular symphysis<\/strong>: Fibrocartilaginous joint between the left and right mandibular segments, located in the midline of the body.<\/p>\n<p class=\"import-Normal\"><strong>Omomyoidea<\/strong>: Order: Primates; Superfamily: Omomyoidea. One of the earliest groups of euprimates (true primates; earliest record in the early Eocene).<\/p>\n<p class=\"import-Normal\"><strong>Petrosal bone<\/strong>: The portion of the temporal bone that houses the inner ear apparatus.<\/p>\n<p class=\"import-Normal\"><strong>Plagiaulacoid<\/strong>: Dental condition where at least one of the lower cheek-teeth (molars or premolars) is a laterally compressed blade.<\/p>\n<p class=\"import-Normal\"><strong>Platyrrhines<\/strong>: Order: Primates; Suborder: Anthropoidea; Infraorder: Platyrrhini. Group containing monkeys found in the Americas.<\/p>\n<p class=\"import-Normal\"><strong>Plesiadapiforms<\/strong>: Order: Plesiadapiformes. Archaic primates or primate-like placental mammals (Early Paleocene\u2013Late Eocene).<\/p>\n<p class=\"import-Normal\"><strong>P<\/strong><strong>lesiomorphic<\/strong>: Having features that are shared by different groups which arose from a common ancestor.<\/p>\n<p class=\"import-Normal\"><strong>Stem<\/strong>: Taxa that are basal to a given crown group but are more closely related to the crown group than to the closest living sister taxon of the crown group.<\/p>\n<p class=\"import-Normal\"><strong>Strepsirrhines<\/strong>: Order: Primates; Suborder: Stresirrhini. Group containing lemurs, lorises, and galagos (does not include tarsiers).<\/p>\n<p class=\"import-Normal\"><strong>Toothcomb<\/strong>: Dental condition found in modern strepsirrhines in which the lower incisors and canines are laterally compressed and protrude forward at a nearly horizontal inclination. This structure is used in grooming.<\/p>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<h2 class=\"import-Normal\">For Further Exploration<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\">Beard, Chris. 2004. <em>The Hunt for the Dawn Monkey: Unearthing the Origins of Monkeys, Apes, and Humans<\/em>. Berkeley: University of California Press.<\/p>\n<p class=\"import-Normal\">Begun, David R. 2010. \u201cMiocene Hominids and the Origins of the African Apes and Humans.\u201d <em>Annual Review of Anthropology<\/em> 39: 67\u201384.<\/p>\n<p class=\"import-Normal\">Fleagle, John G. 2013. <em>Primate Adaptation and Evolution.<\/em> Third edition. San Diego, CA: Academic Press.<\/p>\n<p class=\"import-Normal\">Gebo, Daniel L., ed. 1993. <em>Postcranial Adaptations in Nonhuman Primates<\/em>. Dekalb: Northern Illinois University Press.<\/p>\n<p class=\"import-Normal\">Godfrey, Laurie R., and William L. Jungers. 2002. \u201cQuaternary Fossil Lemurs.\u201d In <em>The Primate Fossil Record, <\/em>edited by Walter C. Hartwig, 97\u2013121. Cambridge: Cambridge University Press.<\/p>\n<p class=\"import-Normal\">Godinot, Marc. 2006. \u201cLemuriform Origins as Viewed from the Fossil Record.\u201d <em>Folia Primatologica<\/em> 77 (6): 446\u2013464.<\/p>\n<p class=\"import-Normal\">Kay, Richard F. 2018. \u201c100 Years of Primate Paleontology.\u201d <em>American Journal of Physical Anthropology<\/em> 165 (4): 652\u2013676.<\/p>\n<p class=\"import-Normal\">Marivaux, Laurent. 2006. \u201cThe Eosimiid and Amphipithecid Primates (Anthropoidea) from the Oligocene of the Bugti Hills (Balochistan, Pakistan): New Insight into Early Higher Primate Evolution in South Asia.\u201d <em>Palaeovertebrata, Montpellier <\/em>34 (1\u20132): 29\u2013109.<\/p>\n<p class=\"import-Normal\">Martin, R. D. 1990. <em>Primate Origins and Evolution<\/em><em>: A <\/em><em>Phylogenetic Reconstruction<\/em>. Princeton: Princeton University Press.<\/p>\n<p class=\"import-Normal\">Rose, Kenneth D., Marc Godinot, and Thomas M. Bown. 1994. \u201cThe Early Radiation of Euprimates and the Initial Diversification of Omomyidae.\u201d In <em>Anthropoid Origins: The Fossil Evidence, <\/em>edited by John G. Fleagle and Richard F. Kay, 1\u201328. New York: Plenum Press.<\/p>\n<p class=\"import-Normal\">Ross, Callum F. 1999. \u201cHow to Carry Out Functional Morphology.\u201d <em>Evolutionary Anthropology<\/em> 7 (6): 217\u2013222.<\/p>\n<p class=\"import-Normal\">Seiffert, Erik R. 2012. \u201cEarly Primate Evolution in Afro-Arabia.\u201d Evolutionary Anthropology: Issues, News, and Reviews 21(6): 239\u2013253.<\/p>\n<p class=\"import-Normal\">Szalay, Frederic S., and Eric Delson. 1979. Evolutionary History of the Primates. New York: Academic Press.<\/p>\n<p class=\"import-Normal\">Ungar, Peter S. 2002. \u201cReconstructing the Diets of Fossil Primates.\u201d In <em>Reconstructing Behavior in the Primate Fossil Record<\/em>, edited by Joseph Plavcan, Richard F. Kay, William Jungers, and Carel P. van Schaik, 261\u2013296. New York: Kluwer Academic\/Plenum Publishers.<\/p>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\">Agust\u00ed, J., A. Sanz de Siria, and M. Garc\u00e9s M. 2003. \u201cExplaining the End of the Hominoid Experiment in Europe.\u201d <em>Journal of Human Evolution<\/em> 45 (2): 145\u2013153.<\/p>\n<p class=\"import-Normal\">Alba, David M., Sergio Alm\u00e9cija, Daniel DeMiguel, Josep Fortuny, Miriam P\u00e9rez de los R\u00edos, Marta Pina, Josep M. 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Simons. 2004. \u201cPhylogenetic, Biogeographic, and Adaptive Implications of New Fossil Evidence Bearing on Crown Anthropoid Origins and Early Stem Catarrhine Evolution.\u201d In <em>Anthropoid Origins: New Visions<\/em>, edited by Callum F. Ross and Richard F. Kay, 157\u2013182. New York: Kluwer\/Plenum Publishing.<\/p>\n<p class=\"import-Normal\">Simons, Elwyn L. 1961. \u201cThe Phyletic Position of <em>Ramapithecus<\/em>.\u201d <em>Postilla<\/em> 57: 1\u20139.<\/p>\n<p class=\"import-Normal\">Simons, Elwyn L. 2001. \u201cThe Cranium of <em>Parapithecus grangeri<\/em>, an Egyptian Oligocene Anthropoidean Primate.\u201d <em>Proceedings of the National Academy of Sciences of the United States of America<\/em> 98 (4): 7892\u20137897.<\/p>\n<p class=\"import-Normal\">Simons, Elwyn L. 2004. \u201cThe Cranium and Adaptations of <em>Parapithecus grangeri<\/em>, a Stem Anthropoid From the Fayum Oligocene of Egypt.\u201d In <em>Anthropoid Origins: New Visions<\/em>, edited by Callum F. Ross and Richard F. Kay, 183\u2013204. New York: Kluwer\/Plenum Publishing.<\/p>\n<p class=\"import-Normal\">Simons, Elwyn L. 2008. \u201cEocene and Oligocene Mammals of the Fayum, Egypt.\u201d In <em>Elwyn Simons: A Search for Origins<\/em>, edited by John G. Fleagle and Christopher C. Gilbert, 87\u2013105. New York: Springer.<\/p>\n<p class=\"import-Normal\">Simons, Elwyn L., and D. Tab Rasmussen. 1994a. \u201cA Remarkable Cranium of <em>Plesiopithecus teras<\/em> (Primates, Prosimii) from the Eocene of Egypt.\u201d <em>Proceedings of the National Academy of Sciences<\/em> <em>of the United States of America<\/em> 91(21): 9946\u20139950.<\/p>\n<p class=\"import-Normal\">Simons, Elwyn L., and D. Tab Rasmussen. 1994b. \u201cA Whole New World of Ancestors: Eocene Anthropoideans from Africa.\u201d <em>Evolutionary Anthropology<\/em> 3 (4): 128\u2013139.<\/p>\n<p class=\"import-Normal\">Simons, Elwyn L., and D. Tab Rasmussen. 1996. \u201cSkull of <em>Catopithecus browni<\/em>, an Early Tertiary Catarrhine.\u201d <em>American Journal of Physical Anthropology<\/em> 100 (2): 261\u2013292.<\/p>\n<p class=\"import-Normal\">Simons, Elwyn L., and Erik R. Seiffert. 1999. \u201cA Partial Skeleton of <em>Proteopithecus<\/em> <em>sylviae<\/em> (Primates Anthropoidea): First Associated Dental and Postcranial Remains of an Eocene Anthropoidean.\u201d <em>Comptes Rendus de l'Acad\u00e9mie des Sciences, Paris<\/em> 329 (12): 921\u2013927.<\/p>\n<p class=\"import-Normal\">Simons, Elwyn L., Erik R. Seiffert, Timothy M. Ryan, and Yousry Attia. 2007. \u201cA Remarkable Female Cranium of the Early Oligocene Anthropoid <em>Aegyptopithecus zeuxis<\/em> (Catarrhini, Propliopithecidae).\u201d <em>Proceedings of the National Academy of Sciences of the United States of America<\/em> 104 (21): 8731\u20138736.<\/p>\n<p class=\"import-Normal\">Simpson, George Gaylord. 1933. \u201cThe \u2018Plagiaulacoid\u2019 Type of Mammalian Dentition: A Study of Convergence.\u201d <em>Journal of Mammalogy<\/em> 14 (2): 97\u2013107.<\/p>\n<p class=\"import-Normal\">Simpson, George Gaylord. 1940. \u201cReview of the Mammal-Bearing Tertiary of South America.\u201d <em>Proceedings of the American Philosophical Society<\/em> 83 (5): 649\u2013709.<\/p>\n<p class=\"import-Normal\">Simpson, George Gaylord. 1967. \u201cThe Tertiary Lorisiform Primates of Africa.\u201d <em>Bulletin of the Museum of Comparative Zoology at Harvard University<\/em> 136: 39\u201362.<\/p>\n<p class=\"import-Normal\">Smith, G. Elliot. 1912. \u201cThe Evolution of Man.\u201d <em>Smithsonian Institute Annual Report <\/em>2012: 553\u2013572.<\/p>\n<p class=\"import-Normal\">Smith, Thierry, Kenneth D. Rose, and Philip D. Gingerich. 2006. \u201cRapid Asia\u2013Europe\u2013North America Geographic Dispersal of Earliest Eocene Primate <em>Teilhardina<\/em> during the Paleocene\u2013Eocene Thermal Maximum.\u201d <em>Proceedings of the National Academy of Sciences of the United States of America<\/em> 103 (30): 11223\u201311227.<\/p>\n<p class=\"import-Normal\">Stehlin, Hans G. 1912. \u201cDie s\u00e4ugetiere des schweizerischen Eocaens. Siebenter teil, erst h\u00e4lfte: <em>Adapis<\/em>\u201d [\u201cThe Mammals of the Swiss Eocene. Part Seven, First Half: Adapis\u201d]. <em>Abhandlungen der Schweizerischen Pal\u00e4ontologischen Gesellschaft<\/em> 38: 1165\u20131298.<\/p>\n<p class=\"import-Normal\">Strait, Suzanne G. 2001. \u201cDietary Reconstruction of Small-Bodied Omomyoid Primates.\u201d <em>Journal of Vertebrate Paleontology<\/em> 21 (2): 322\u2013334.<\/p>\n<p class=\"import-Normal\">Sussman, Robert W. 1991. \u201cPrimate Origins and the Evolution of Angiosperms.\u201d <em>American Journal of Primatology<\/em> 23 (4): 209\u2013223.<\/p>\n<p class=\"import-Normal\">Suwa, Gen, Reiko T. Kono, Shigehiro Katoh, Berhane Asfaw, and Yonas Beyene. 2007. \u201cA New Species of Great Ape from the Late Miocene Epoch in Ethiopia.\u201d <em>Nature<\/em> 448 (7156): 921\u2013924.<\/p>\n<p class=\"import-Normal\">Teaford, Mark F., Mary C. Maas, and Elwyn L. Simons. 1996. \u201cDental Microwear and Microstructure in Early Oligocene Primates from the Fayum, Egypt: Implications for Diet.\u201d <em>American Journal of Physical Anthropology<\/em> 101 (4): 527\u2013543.<\/p>\n<p class=\"import-Normal\">Ungar, Peter S., and Richard F. Kay. 1995. \u201cThe Dietary Adaptations of European Miocene Catarrhines.\u201d <em>Proceedings of the National Academy of Sciences of the United States of America<\/em> 92 (12): 5479\u20135481.<\/p>\n<p>Ungar, Peter S. 2002. \u201cReconstructing the Diets of Fossil Primates.\u201d In Reconstructing Behaviour in the Primate Fossil Record, edited by Joseph Plavcan, Richard F. Kay, William Jungers, and Carel P. van Schaik, 261\u2013296. New York: Kluwer Academic\/Plenum Publishers.<\/p>\n<p class=\"import-Normal\">Wang, Cui-Bin, Ling-Xia Zhao, Chang-Zhu Jin, Yuan Wang, Da-Gong Qin, and Wen-Shi Pan. 2014. \u201cNew Discovery of Early Pleistocene Orangutan Fossils from Sanhe Cave in Chongzuo, Guangxi, Southern China.\u201d <em>Quaternary International<\/em> 354: 68\u201374.<\/p>\n<p class=\"import-Normal\">Ward, C. V., A. Walker, and M. F. Teaford. 1991. \u201c<em>Proconsul<\/em> Did Not Have a Tail.\u201d <em>Journal of Human Evolution<\/em> 21 (3): 215\u2013220.<\/p>\n<p class=\"import-Normal\">Wheeler, Brandon C. 2010. \u201cCommunity Ecology of the Middle Miocene Primates of La Venta, Colombia: The Relationship between Ecological Diversity, Divergence Time, and Phylogenetic Richness.\u201d <em>Primates<\/em> 51 (2): 131\u2013138.<\/p>\n<p class=\"import-Normal\">Williams, Blythe A., and Richard F. Kay. 1995. \u201cThe Taxon Anthropoidea and the Crown Clade Concept.\u201d <em>Evolutionary Anthropology<\/em> 3 (6): 188\u2013190.<\/p>\n<p class=\"import-Normal\">Williams, Blythe A., Richard F. Kay, and E. Christopher Kirk. 2010a. \u201cNew Perspectives on Anthropoid Origins.\u201d <em>Proceedings of the National Academy<\/em> <em>of the United States of America<\/em> 107 (11): 4797\u20134804.<\/p>\n<p class=\"import-Normal\">Williams, Blythe A., Richard F. Kay, E. Christopher Kirk, and Callum F. Ross. 2010b. \u201c<em>Darwinius masillae<\/em> Is a European Middle Eocene Stem Strepsirrhine\u2014A Reply to Franzen et al.\u201d <em>Journal of Human Evolution<\/em> 59(5): 567\u2013573.<\/p>\n<p class=\"import-Normal\">Wilson Mantilla, G. P., S. G. B. Chester, W. A. Clemens, J. R. Moore, C. J. Sprain, B. T. Hovatter, W. S. Mitchell, W. W. Mans, R. Mundil, and P. R. Renne. 2021. \u201cEarliest Palaeocene Purgatoriids and the Initial Radiation of Stem Primates.\u201d <em>Royal Society Open Science<\/em> 8(2):210050. doi:10.1098\/rsos.210050.<\/p>\n<h2 class=\"import-Normal\">Acknowledgements<\/h2>\n<p class=\"import-Normal\">We are immensely grateful to the editors of this book, Drs. Beth Shook, Lara Braff, Katie Nelson, and Kelsie Aguilera, for their time and commitment to making this knowledge freely accessible to all, and for giving us the opportunity to participate in this important project.<\/p>\n<\/div>\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_1683_1684\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1684\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\">Jonathan M. G. Perry, Ph.D., Western University of Health Sciences<\/p>\n<p class=\"import-Normal\">Stephanie L. Canington, Ph.D., University of Pennsylvania<\/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__-10\/\"><em>Chapter 8: Primate Evolution<\/em><\/a><em>\u201d by Jonathan M. G. Perry and Stephanie L. Canington. 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\">Learning Objectives<\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li>Understand the major trends in primate evolution from the origin of primates to the origin of our own species.<\/li>\n<li>Learn about primate adaptations and how they characterize major primate groups.<\/li>\n<li>Discuss the kinds of evidence that anthropologists use to find out how extinct primates are related to each other and to living primates.<\/li>\n<li>Recognize how the changing geography and climate of Earth have influenced where and when primates have thrived or gone extinct.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p class=\"import-Normal\">The first fifty million years of primate evolution was a series of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1683\">adaptive radiations<\/a><\/strong> leading to the diversification of the earliest lemurs, monkeys, and apes. The primate story begins in the canopy and understory of conifer-dominated forests, with our small, furtive ancestors subsisting at night, beneath the notice of day-active dinosaurs.<\/p>\n<p class=\"import-Normal\">From the ancient <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1684\"><strong>plesiadapiforms<\/strong><\/a> (archaic primates) to the earliest groups of true primates (<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1686\"><strong>euprimates<\/strong><\/a>) (Bloch and Boyer 2002), the origin of our own order is characterized by the struggle for new food sources and microhabitats in the arboreal setting. Climate change forced major extinctions as the northern continents became increasingly dry, cold, and seasonal and as tropical rainforests gave way to deciduous forests, woodlands, and eventually grasslands. Lemurs, lorises, and tarsiers\u2014once diverse groups containing many species\u2014became rare, except for lemurs in Madagascar, where there were no anthropoid competitors and perhaps few predators. Meanwhile, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1685\">anthropoids<\/a><\/strong> (monkeys and apes) likely emerged in Asia and then dispersed across parts of the northern hemisphere, Africa, and ultimately South America. The movement of continents, shifting sea levels, and changing patterns of rainfall and vegetation contributed to the developing landscape of primate biogeography, morphology, and behaviour. Today\u2019s primates provide modest reminders of the past diversity and remarkable adaptations of their extinct relatives. This chapter explores the major trends in primate evolution from the origin of the Order Primates to the beginnings of our own lineage, providing a window into these stories from our ancient past.<\/p>\n<h2 class=\"import-Normal\">Major Hypotheses About Primate Origins<\/h2>\n<p class=\"import-Normal\">For many groups of mammals, there is a key feature that led to their success. A good example is powered flight in bats. Primates lack a feature like this (see Chapter 6). Instead, if there is something unique about primates, it is probably a group of features rather than one single thing. Because of this, anthropologists and palaeontologists struggle to describe an ecological scenario that could explain the rise and success of our own order. Three major hypotheses have been advanced to consider the origin of primates and to explain what makes our order distinct among mammals (Figure 9.1); these are described below.<\/p>\n<figure id=\"attachment_277-11\" aria-describedby=\"caption-attachment-277-11\" style=\"width: 634px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-255\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2023\/03\/8.1.jpg\" alt=\"Primates swinging in tree, eating an insect, and eating fruit.\" width=\"634\" height=\"221\" \/><figcaption id=\"caption-attachment-277-11\" class=\"wp-caption-text\">Figure 9.1: The three major hypotheses are (a) the arboreal hypothesis, (b) the visual predation hypothesis, and (c) the angiosperm-primate coevolution hypothesis. Credit: Primate origin hypotheses original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by <a class=\"rId13\" href=\"https:\/\/marynelsonstudio.com\">Mary Nelson<\/a> is under a <a class=\"rId14\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Arboreal Hypothesis<\/strong><\/h3>\n<p class=\"import-Normal\">In the 1800s, many anthropologists viewed all animals in relation to humans. That is, animals that were more like humans were considered to be more \u201cadvanced\u201d and those lacking humanlike features were considered more \u201cprimitive.\u201d This way of thinking was particularly obvious in studies of primates. A more modern way of referring to members of a group that lack certain evolutionary innovations seen in other members is to call them <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1688\"><strong>plesiomorphic<\/strong><\/a> (literally \u201canciently shaped\u201d). The state of their morphological features is sometimes referred to as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1689\"><strong>ancestral<\/strong><strong> traits<\/strong><\/a>.<\/p>\n<p class=\"import-Normal\">Thus, when anthropologists sought features that separate primates from other mammals, they focused on features that were least developed in lemurs and lorises, more developed in monkeys, and most developed in apes (Figure 9.2). Frederic Wood Jones, one of the leading anatomist-anthropologists of the early 1900s, is usually credited with the Arboreal Hypothesis of primate origins (Jones 1916). This hypothesis holds that many of the features of primates evolved to improve locomotion in the trees; this way of getting around is referred to as arboreal. For example, the grasping hands and feet of primates are well suited to gripping tree branches of various sizes and our flexible joints are good for reorienting the extremities in many different ways. A mentor of Jones, Grafton Elliot Smith, had suggested that the reduced olfactory system, acute vision, and forward-facing eyes of primates are adaptations for making accurate leaps and bounds through a complex, three-dimensional canopy (Smith 1912). The forward orientation of the eyes in primates causes the visual fields to overlap, enhancing depth perception, especially at close range. Evidence to support this hypothesis includes the facts that many extant primates are arboreal, and the plesiomorphic members of most primate groups are dedicated arborealists. The Arboreal Hypothesis was well accepted by most anthropologists at the time and for decades afterward.<\/p>\n<figure style=\"width: 663px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image27-2.png\" alt=\"Diagram shows primates descended from Plesiadapiforms.\" width=\"663\" height=\"543\" \/><figcaption class=\"wp-caption-text\">Figure 9.2: Primate family tree showing major groups. Disconnected lines show uncertainty about relationships. Two lines lead to tarsiers from different possible groups of origin. <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 class=\"rId16\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Primate family tree (Figure 8.2)<\/a> by Jonathan M. G. Perry 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<h3 class=\"import-Normal\"><strong>Visual Predation Hypothesis<\/strong><\/h3>\n<p class=\"import-Normal\">In the late 1960s and early 1970s, Matt Cartmill studied and tested the idea that the characteristic features of primates evolved in the context of arboreal locomotion. Cartmill noted that squirrels climb trees (and even vertical walls) very effectively, even though they lack some of the key adaptations of primates. As members of the Order Rodentia, squirrels also lack the hand and foot anatomy of primates. They have claws instead of flattened nails and their eyes face more laterally than those of primates. Cartmill reasoned that there must be some other explanation for the unique traits of primates. He noted that some nonarboreal animals share at least some of these traits with primates; for example, cats and predatory birds have forward-facing eyes that enable visual field overlap. Cartmill suggested that the unique suite of features in primates is an adaptation to detecting insect prey and guiding the hands (or feet) to catch insects (Cartmill 1972). His hypothesis emphasizes the primary role of vision in prey detection and capture; it is explicitly comparative, relying on form-function relationships in other mammals and nonmammalian vertebrates. According to Cartmill, many of the key features of primates evolved for preying on insects in this special manner (Cartmill 1974).<\/p>\n<h3 class=\"import-Normal\"><strong>Angiosperm-Primate Coevolution Hypothesis<\/strong><\/h3>\n<p class=\"import-Normal\">The visual predation hypothesis was unpopular with some anthropologists. One reason for this is that many primates today are not especially predatory. Another is that, whereas primates do seem well adapted to moving around in the smallest, terminal branches of trees, insects are not necessarily easier to find there. A counterargument to the visual predation hypothesis is the angiosperm-primate coevolution hypothesis. Primate ecologist Robert Sussman (Sussman 1991) argued that the few primates that eat mostly insects often catch their prey on the ground rather than in tree branches. Furthermore, predatory primates often use their ears more than their eyes to detect prey. Finally, most early primate fossils show signs of having been omnivorous rather than insectivorous. Instead, he argued, the earliest primates were probably seeking fruit. Fruit (and flowers) of angiosperms (flowering plants) often develop in the terminal branches. Therefore, any mammal trying to access those fruits must possess anatomical traits that allow them to maintain their hold on thin branches and avoid falling while reaching for the fruits. Primates likely evolved their distinctive visual traits and extremities in the Paleocene (approximately 65 million to 54 million years ago) and Eocene (approximately 54 million to 34 million years ago) epochs, just when angiosperms were going through a revolution of their own\u2014the evolution of large, fleshy fruit that would have been attractive to a small arboreal mammal. Sussman argued that, just as primates were evolving anatomical traits that made them more efficient fruit foragers, angiosperms were also evolving fruit that would be more attractive to primates to promote better seed dispersal. This mutually beneficial relationship between the angiosperms and the primates was termed coevolution or more specifically <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1691\"><strong>diffuse coevolution<\/strong>.<\/a><\/p>\n<p class=\"import-Normal\">At about the same time, D. Tab Rasmussen noted several parallel traits in primates and the South American woolly opossum, <em>Caluromys<\/em>. He argued that early primates were probably foraging on both fruits and insects (Rasmussen 1990). As is true of <em>Caluromys<\/em> today, early primates probably foraged for fruits in the terminal branches of angiosperms, and they probably used their visual sense to aid in catching insects. Insects are also attracted to fruit (and flowers), so these insects represent a convenient opportunity for a primarily fruit-eating primate to gather protein. This solution is a compromise between the visual predation hypothesis and the angiosperm-primate coevolution hypothesis. It is worth noting that other models of primate origins have been proposed, and these include the possibility that no single ecological scenario can account for the origin of primates.<\/p>\n<h2 class=\"import-Normal\">The Origins of Primates<\/h2>\n<h3 class=\"import-Normal\"><strong>Paleocene: Mammals in the Wake of Dinosaur Extinctions<\/strong><\/h3>\n<p class=\"import-Normal\">Placental mammals, including primates, originated in the Mesozoic Era (approximately 251 million to 65.5 million years ago), the Age of Dinosaurs. During this time, most placental mammals were small, probably nocturnal, and probably avoided predators via camouflage and slow, quiet movement. It has been suggested that the success and diversity of the dinosaurs constituted a kind of ecological barrier to Mesozoic mammals. The extinction of the dinosaurs (and many other organisms) at the end of the Cretaceous Period (approximately 145.5\u201365.5 million years ago) might have opened up these ecological niches, leading to the increased diversity and disparity in mammals of the Tertiary Period (approximately 65.5\u20132.5 million years ago).<\/p>\n<p class=\"import-Normal\">The Paleocene was the first epoch in the Age of Mammals. Soon after the Cretaceous-Tertiary (K-T) extinction event, new groups of placental mammals appear in the fossil record. Many of these groups achieved a broad range of sizes and lifestyles as well as a great number of species before declining sometime in the Eocene (or soon thereafter). These groups were ultimately replaced by the modern orders of placental mammals (Figure 9.3). It is unknown whether these replacements occurred gradually, for example by competitive exclusion, or rapidly, perhaps by sudden geographic dispersals with replacement. In some senses, the Paleocene might have been a time of recovery from the extinction event; it was cooler and more seasonal globally than the subsequent Eocene.<\/p>\n<figure style=\"width: 628px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image26.jpg\" alt=\"Person in front of a mural depicting forest animals.\" width=\"628\" height=\"511\" \/><figcaption class=\"wp-caption-text\">Figure 9.3: A mural of Eocene flora and fauna in North America. Credit: <a class=\"rId19\" href=\"https:\/\/flickr.com\/photos\/126377022@N07\/18404106406\">Image from page 27 of \"Annual report for the year ended June 30 ...\" (1951)<\/a> by <a class=\"rId20\" href=\"https:\/\/flickr.com\/photos\/internetarchivebookimages\/\">Internet Archive Book Images<\/a> has been designated to the <a class=\"rId21\" href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/\">public domain (CC0)<\/a>. This photograph of the mural \"Fauna and flora of middle Eocene in the Wyoming area\" by Jay Matternes, was originally published by the <a class=\"rId22\" href=\"https:\/\/www.si.edu\/\">Smithsonian<\/a>, and can be viewed in context in the <a class=\"rId23\" href=\"https:\/\/archive.org\/details\/annualreportfory1961united\/page\/7\/mode\/1up?view=theater\">online version of this book<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Plesiadapiforms, the Archaic Primates<\/strong><\/h3>\n<p class=\"import-Normal\">The Paleocene epoch saw the emergence of several families of mammals that have been implicated in the origin of primates. These are the plesiadapiforms, which are archaic primates, meaning they possessed some primate features and lacked others. The word <em>plesiadapiform <\/em>means \u201calmost adapiform,\u201d a reference to some similarities between some plesiadapiforms and some adapiforms (or adapoids; later-appearing true primates)\u2014mainly in the molar teeth. Because enamel fossilizes better than other parts of the body, the molar teeth are the parts most often found and first discovered for any new species. Thus, dental similarities were often the first to be noticed by early mammalian paleontologists, partly explaining why plesiadapiforms were thought to be primates. Major morphological differences between plesidapiforms and euprimates (true primates) were observed later when more parts of plesiadapiform skeletons were discovered. Many plesiadapiforms have unusual anterior teeth and most have digits possessing claws rather than nails. So far, no plesiadapiform ever discovered has a postorbital bar (seen in extant <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1712\">strepsirrhines<\/a><\/strong>) or septum (as seen in <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1713\">haplorhines<\/a><\/strong>), and whether or not the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1711\">auditory bulla<\/a><\/strong> was formed by the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1714\"><strong>petrosal bone<\/strong> <\/a>remains unclear for many plesiadapiform specimens. Nevertheless, there are compelling reasons (partly from new skeletal material) for including plesiadapiforms within the Order Primates.<\/p>\n<h4 class=\"import-Normal\"><em>Geographic and Temporal Distribution<\/em><\/h4>\n<p class=\"import-Normal\"><em>Purgatorius<\/em> is generally considered to be the earliest primate. This Paleocene mammal is known from teeth that are very plesiomorphic for a primate. It has some characteristics that suggest it is a basal plesiadapiform, but there is very little to link it specifically with euprimates (see Clemens 2004). Its ankle bones suggest a high degree of mobility, signaling an arboreal lifestyle (Chester Et al. 2015). <em>Purgatorius<\/em> is plesiomorphic enough to have given rise to all primates, including the plesiadapiforms. However, new finds suggest that this genus was more diverse and had more differing tooth morphologies than previously appreciated (Wilson Mantilla Et al. 2021). Plesiadapiform families were numerous and diverse during parts of the Paleocene in western North America and western Europe, with some genera (e.g., <em>Plesiadapis<\/em>; see Figure 9.4) living on both continents (Figure 9.5). Thus, there were probably corridors for plesiadapiform dispersal between the two continents, and it stands to reason that these mammals were living all across North America, including in the eastern half of the continent and at high latitudes. A few plesiadapiforms have been described from Asia (e.g., <em>Carpocristes<\/em>), but the affinities of these remain uncertain.<\/p>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 473.25pt\">\n<caption>Figure 9.4: Families of plesiadapiforms with example genera and traits: a table. Credit: Plesiadapiforms table original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Jonathan M. G. Perry and Stephanie L. Canington is under a <a class=\"rId24\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Content derived from Fleagle 2013.<\/caption>\n<thead>\n<tr style=\"height: 25pt\">\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Family<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Genera<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Morphology<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Location<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Age<\/strong><sup><strong>1<\/strong><\/sup><\/p>\n<p>&nbsp;<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table1-R\" style=\"height: 17pt\">\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Paromomyidae<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Ignacius<\/em><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Long, dagger-like, lower incisor.<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">North America and Europe<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Early Paleocene to Late Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 18pt\">\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Carpolestidae<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Carpolestes<\/em><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Plagiaulacoid dentition. Limb adaptations to terminal branch feeding. Grasping big toe.<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">North America, Europe, and Asia<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Middle Paleocene to Early Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 16pt\">\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Plesiadapidae<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Plesiadapis<\/em><\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Mitten-like upper incisor. Diastema. Large body size for group.<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">North America and Europe<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Middle Paleocene to Early Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table1-R\" style=\"height: 1pt\">\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\" colspan=\"4\">\n<p class=\"import-Normal\"><sup>1<\/sup> Derived from Fleagle 2013.<\/p>\n<\/td>\n<td class=\"Table1-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\">\n<p class=\"import-Normal\">\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<figure id=\"attachment_277-12\" aria-describedby=\"caption-attachment-277-12\" style=\"width: 555px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-258 \" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5-5-e1691791897574.png\" alt=\"Global map with not fully formed continents.\" width=\"555\" height=\"308\" \/><figcaption id=\"caption-attachment-277-12\" class=\"wp-caption-text\">Figure 9.5: Map of the world in the Paleocene, highlighting plesiadapiform localities on lands that would become North America, southern Europe, and eastern Asia. Credit: <a class=\"rId26\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Paleocene Map with Plesiadapiform Localities (Figure 8.4)<\/a> original to<a class=\"rId27\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"> Expl<\/a><a class=\"rId28\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">orations: An Open Invitation to Biological Anthropology<\/a> by Elyssa Ebding at <a class=\"rId29\" href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\">GeoPlace, California State University, Chico<\/a> is under a <a class=\"rId30\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Localities based on Fleagle 2013, 211.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>General Morphological Features<\/em><\/h4>\n<p class=\"import-Normal\">Although there is much morphological variation among the families of plesiadapiforms, some common features unite the group. Most plesiadapiforms were small, the largest being about three kilograms (approximately 7 lbs.; <em>Plesiadapis cookei<\/em>). They had small brains and fairly large snouts, with eyes that faced more laterally than in euprimates. Many species show reduction and\/or loss of the canine and anterior premolars, with the resulting formation of a rodent-like <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1172\">diastema<\/a> <\/strong>(a pronounced gap between the premolars and the incisors, with loss of at least the canine); this probably implies a herbivorous diet. Some families appear to have had very specialized diets, as suggested by unusual tooth and jaw shapes.<\/p>\n<p class=\"import-Normal\">Arguably the most interesting and unusual family of plesiadapiforms is the Carpolestidae. They are almost exclusively from North America (with a couple of possible members from Asia), and mainly from the Middle and Late Paleocene. Their molars are not very remarkable, being quite similar to those of some other plesiadapiforms (e.g., Plesiadapidae). However, their lower posterior premolars (p4) are laterally compressed and blade-like with vertical serrations topped by tiny cuspules. This unusual dental morphology is termed <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1693\"><strong><em>plagiaulacoid<\/em><\/strong> <\/a> (Simpson 1933). The upper premolar occlusal surfaces are broad and are covered with many small cuspules; the blade-like lower premolar might have cut across these cuspules, between them, or both.<\/p>\n<div class=\"mceTemp\"><\/div>\n<figure style=\"width: 314px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image1-5.png\" alt=\"Small brown animal with long tail.\" width=\"314\" height=\"178\" \/><figcaption class=\"wp-caption-text\">Figure 9.6: An artistic rendition of Carpolestes simpsoni moving along a small diameter support. Credit: <a class=\"rId32\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:CarpolestesCL.png\">CarpolestesCL<\/a> by <a class=\"rId33\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Sisyphos23\">Sisyphos23<\/a> is under a <a class=\"rId34\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Many plesiadapiforms have robust limb bones with hallmarks of arboreality. Instead of having nails, most taxa had sharp claws on most or all of the digits. The extremities show grasping abilities comparable to those of primates and some arboreal marsupials. Nearly complete skeletons have yielded a tremendous wealth of information on locomotor and foraging habits. Many plesiadapiforms appear to have been able to cling to vertical substrates (like a broad tree trunk) using their sharp claws, propelling themselves upward using powerful hindlimbs, bounding along horizontal supports, grasping smaller branches, and moving head-first down tree trunks. In carpolestids in particular, the skeleton appears to have been especially well adapted to moving slowly and carefully in small terminal branches (Figure 9.6).<\/p>\n<\/div>\n<div class=\"textbox shaded\">\n<h3 class=\"import-Normal\">Dig Deeper: Debate on the Relationship of Plesiadapiforms to True Primates<\/h3>\n<p class=\"import-Normal\">In the middle of the twentieth century, treeshrews (Order Scandentia) were often considered part of the Order Primates, based on anatomical similarities between some treeshrews and primates. For many people, plesiadapiforms represented intermediates between primates and treeshrews, so plesiadapiforms were included in Primates as well.<\/p>\n<p class=\"import-Normal\">Studies of reproduction and brain anatomy in treeshrews and lemurs suggested that treeshrews are not primates (e.g., Martin 1968). This was soon followed by the suggestion to also expel plesiadapiforms (Martin 1972) from the Order Primates. Like treeshrews, plesiadapiforms lack a postorbital bar, nails, and details of the ear region that characterize true primates. Many paleoanthropologists were reluctant to accept this move to banish plesiadapiforms (e.g., F. S. Szalay, P. D. Gingerich).<\/p>\n<p class=\"import-Normal\">Later, K. Christopher Beard (1990) found that in some ways, the digits of paromomyid plesiadapiforms are actually more similar to those of dermopterans (Order Dermoptera), the closest living relatives of primates, than they are to those of primates themselves (but see Krause 1991). At the same time, Richard Kay and colleagues (1990) found that cranial circulation patterns and auditory bulla morphology in the paromomyid, <em>Ignacius <\/em>(see Figure 9.4), are more like those of dermopterans than of primates.<\/p>\n<p class=\"import-Normal\">For many anthropologists, this one-two punch effectively removed plesiadapiforms from the Order Primates. In the last two decades, the tide of opinion has turned again, with many researchers reinstating plesiadapiforms as members of the Order Primates. New and more complete specimens demonstrate that the postcranial skeletons of plesiadapiforms, including the hands and feet, were primate-like, not dermorpteran-like (Bloch and Boyer 2002, 2007). New fine-grained CT scans of relatively complete plesiadapiform skulls revealed that they share some key traits with primates to the exclusion of other placental mammals (Bloch and Silcox 2006). Most significant was the suggestion that <em>Carpolestes simpsoni <\/em>possessed an auditory bulla formed by the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1696\"><strong>petrosal <\/strong><strong>bone<\/strong><\/a>, like in all living primates.<\/p>\n<p class=\"import-Normal\">The debate about the status of plesiadapiforms continues, owing to a persistent lack of key bones in some species and owing to genuine complexity of the anatomical traits involved. Maybe plesiadapiforms were the ancestral stock from which all primates arose, with some plesiadapiforms (e.g., carpolestids) nearer to the primate <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1723\">stem<\/a><\/strong> than others.<\/p>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<h3 class=\"import-Normal\"><strong>Adapoids and Omomyoids, the First True Primates<\/strong><\/h3>\n<h4 class=\"import-Normal\"><em>Geographic and Temporal Distribution<\/em><\/h4>\n<p class=\"import-Normal\">The first universally accepted fossil primates are the adapoids (Superfamily <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1695\"><strong>Adapoidea<\/strong><\/a>) and the omomyoids (Superfamily <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1694\">Omomyoidea<\/a>)<\/strong>. These groups become quite distinct over evolutionary time, filling mutually exclusive niches for the most part. However, the earliest adapoids are very similar to the earliest omomyoids.<\/p>\n<p class=\"import-Normal\">The adapoids were mainly diurnal and herbivorous, with some achieving larger sizes than any plesiadapiforms (10 kg; 22 lbs.). By contrast, the omomyoids were mainly nocturnal, insectivorous and frugivorous, and small.<\/p>\n<p class=\"import-Normal\">Both groups appear suddenly at the start of the Eocene, where they are present in western North America, western Europe, and India (Figure 9.7). This wide dispersal of early primates was probably due to the presence of rainforest corridors extending far into northern latitudes.<\/p>\n<figure id=\"attachment_277-13\" aria-describedby=\"caption-attachment-277-13\" style=\"width: 539px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-260\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image22-3-e1691792023503.png\" alt=\"Global map with not fully formed continents and omomyoid localities.\" width=\"539\" height=\"317\" \/><figcaption id=\"caption-attachment-277-13\" class=\"wp-caption-text\">Figure 9.7: Map of the world in the Eocene, highlighting adapoid and omomyoid localities on lands that would become North America, southern Europe, Africa, and Asia. Credit: <a class=\"rId36\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Eocene Map with Adapoid and Omomyoid Localities (Figure 8.6)<\/a> original to <a class=\"rId37\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Elyssa Ebding at <a class=\"rId38\" href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\">GeoPlace, California State University, Chico<\/a> is under a <a class=\"rId39\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Localities based on Fleagle 2013, 229.<\/figcaption><\/figure>\n<p class=\"import-Normal\">In North America and Europe, both groups achieved considerable diversity in the Middle Eocene, then mostly died out at the end of that epoch (Figure 9.8). In some Eocene rock formations in the western United States, adapoids and omomyoids make up a major part of the mammalian fauna. The Eocene of India has yielded a modest diversity of euprimates, some of which are so plesiomorphic that it is difficult to know whether they are adapoids or omomyoids (or even early anthropoids).<\/p>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 473.25pt\">\n<caption>Figure 9.8: Families of adapoids and omomyoids with example genera and traits: a table. Credit: Adapoids and omomyoids table original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Jonathan M. G. Perry and Stephanie L. Canington is under a <a class=\"rId40\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Content derived from Fleagle 2013.<\/caption>\n<thead>\n<tr style=\"height: 25pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Family<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Genera<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Morphology<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Location<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Age<\/strong><sup><strong>1<\/strong><\/sup><\/p>\n<p>&nbsp;<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table2-R\" style=\"height: 18pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Cercamoniidae<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Donrussellia<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Variable in tooth number and jaw shape.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Europe and Asia<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Early to Late Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 16pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Asiadapidae<sup>2<\/sup><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Asiadapis<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Plesiomorphic teeth and jaw resemble early Omomyids.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Asia<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Early Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 16pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Caenopithecidae<sup>3<\/sup><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Darwinius<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Robust jaws with crested molars. Fewer premolars.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Europe, Africa, North America, and Asia<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Middle to Late Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 16pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Adapidae<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Adapis<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Fused mandible. Long molar crests. Large size and large chewing muscles.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Europe<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Late Eocene to Early Oligocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 16pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Sivaladapidae<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Sivaladapis<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Some large with robust jaws.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Asia<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Middle Eocene to Late Miocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 16pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Notharctidae<sup>4<\/sup><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Notharctus<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Canine sexual dimorphism. Lemur-like skull. Clinging and leaping adaptations.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">North America and Europe<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Early to Middle Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 16pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Omomyidae<sup>5<\/sup><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Teilhardina<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Small, nocturnal, frugivorous or insectivorous. Tarsier-like skull in some.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">North America, Europe, and Asia<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Early Eocene to Early Miocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 16pt\">\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Microchoeridae<sup>6<\/sup><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Necrolemur<\/em><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Long bony ear tubes. Tarsier-like lower limb adaptations for leaping.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Europe and Asia<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Early Eocene to Early Oligocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 1pt\">\n<td class=\"Table2-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\" colspan=\"4\">\n<p class=\"import-Normal\"><sup>1<\/sup> Derived from Fleagle 2013.<\/p>\n<p class=\"import-Normal\"><sup>2<\/sup> See Dunn Et al. 2016 and Rose Et al. 2018.<\/p>\n<p class=\"import-Normal\"><sup>3<\/sup> See Kirk and Williams 2011 and Seiffert Et al. 2009.<\/p>\n<p class=\"import-Normal\"><sup>4<\/sup> See Gregory 1920.<\/p>\n<p class=\"import-Normal\"><sup>5<\/sup> See Beard and MacPhee 1994 and Strait 2001.<\/p>\n<p class=\"import-Normal\"><sup>6<\/sup> See Schmid 1979.<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\">\n<p class=\"import-Normal\">\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<p class=\"import-Normal\">Adapoids and omomyoids barely survived the Eocene-Oligocene extinctions, when colder temperatures, increased seasonality, and the retreat of rainforests to lower latitudes led to changes in mammalian biogeography. In North America, one genus (originally considered an omomyoid but recently reclassified as Adapoidea) persisted until the Miocene: <em>Ekgmowechashala<\/em> (Rose and Rensberger 1983). This taxon has highly unusual teeth and might have been a late immigrant to North America from Asia. In Asia, one family of adapoids, the Sivaladapidae, retained considerable diversity as late as the Late Miocene.<\/p>\n<h4 class=\"import-Normal\"><em>Adapoid Diversity<\/em><\/h4>\n<p class=\"import-Normal\">Adapoids were very diverse, particularly in the Eocene of North America and Europe. They can be divided into six families, with a few species of uncertain familial relationship. As a group, adapoids have some features in common, although much of what they share is plesiomorphic. Important features include the hallmarks of euprimates: postorbital bar, flattened nails, grasping extremities, and a petrosal bulla (Figures 9.9 and 9.10). In addition, some adapoids retain the ancestral dental formula of 2.1.4.3; that is, in each quadrant of the mouth, there are two incisors, one canine, four premolars, and three molars. In general, the incisors are small compared to the molars, but the canines are relatively large, with sexual dimorphism in some species. Cutting crests on the molars are well developed in some species, and the two halves of the mandible were fused at the midline in some species. Some adapoids were quite small (<em>Anchomomys <\/em>at a little over 100 g), and some were quite large (<em>Magnadapis<\/em> at 10 kg; 22 lbs.). Furthermore, the spaces and attachment features for the chewing muscles were truly enormous in some species, suggesting that these muscles were very large and powerful. Taken together, this suggests an overall adaptive profile of diurnal herbivory. The canine sexual dimorphism in some species suggests a possible mating pattern of polygyny, as males in polygynous primate species often compete with each other for mates and have especially large canine teeth.<\/p>\n<figure style=\"width: 548px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image18-1.jpg\" alt=\"Three partial animal crania.\" width=\"548\" height=\"350\" \/><figcaption class=\"wp-caption-text\">Figure 9.9: Representative crania of Adapidae from Museum d\u2019Histoire Naturelle Victor Brun, a natural history museum in Montauban, France. The white scale bar is 1 cm long. Credit: <a class=\"rId43\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Representative crania of adapids (European adapoids, (Figure 8.7)<\/a> from the <a class=\"rId44\" href=\"https:\/\/www.museum.montauban.com\/\">Museum d\u2019Histoire Naturelle Victor Brun in Montauban, France<\/a> original to <a class=\"rId45\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology <\/a>by Jonathan M. G. Perry is under a <a class=\"rId46\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<figure style=\"width: 547px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image19-2.jpg\" alt=\"Side views of small rodentlike skeleton with long tail.\" width=\"547\" height=\"525\" \/><figcaption class=\"wp-caption-text\">Figure 9.10: Darwinius masillae, a member of the Caenopithecidae. The slab on the left is Plate A and the slab on the right is Plate B. The parts of the skeleton in B that are outside of the dashed lines were fabricated. Credit: <a class=\"rId48\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Darwinius%20masillae%20holotype%20slabs.jpg\">Darwinius masillae holotype slabs<\/a> by Jens L. Franzen, Philip D. Gingerich, J\u00f6rg Habersetzer1, J\u00f8rn H. Hurum, Wighart von Koenigswald, B. Holly Smith is under a <a class=\"rId49\" href=\"https:\/\/creativecommons.org\/licenses\/by\/2.5\/legalcode\">CC BY 2.5 License<\/a>. Originally from Franzen Et al. 2009.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Omomyoid Diversity<\/em><\/h4>\n<p class=\"import-Normal\">Like adapoids, omomyoids appeared suddenly at the start of the Eocene and then became very diverse with most species dying out before the Oligocene. Omomyoids are known from thousands of jaws with teeth, relatively complete skulls for about a half-dozen species, and very little postcranial material. Omomyoids were relatively small primates, with the largest being less than three kilograms (approximately 7 lbs.; <em>Macrotarsius montanus<\/em>). All known crania possess a postorbital bar, which in some has been described as \u201cincipient closure.\u201d Some\u2014but not all\u2014known crania have an elongated bony ear tube extending lateral to the location of the eardrum, a feature seen in living tarsiers and <strong>catarrhines<\/strong>. The anterior teeth tend to be large, with canines that are usually not much larger than the incisors. Often it is difficult to distinguish closely related species using molar morphology, but the premolars tend to be distinct from one species to another. The postcranial skeleton of most omomyoids shows hallmarks of leaping behaviour reminiscent of that of tarsiers. In North America, omomyoids became very diverse and abundant. In fact, omomyoids from Wyoming are sufficiently abundant and from such stratigraphically controlled conditions that they have served as strong evidence for the gradual evolution of anatomical traits over time (Rose and Bown 1984).<\/p>\n<p class=\"import-Normal\"><em>Teilhardina <\/em>(Figure 9.11; see Figure 9.2) is one of the earliest and arguably the most plesiomorphic of omomyoids. <em>Teilhardina<\/em> has several species, most of which are from North America, with one from Europe (<em>T. belgica<\/em>) and one from Asia (<em>T. asiatica<\/em>). The species of this genus are anatomically similar and the deposits from which they are derived are roughly contemporaneous. Thus, this small primate likely dispersed across the northern continents very rapidly (Smith Et al. 2006).<\/p>\n<figure style=\"width: 545px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image3-1.jpg\" alt=\"World map with primates jumping across forested areas.\" width=\"545\" height=\"289\" \/><figcaption class=\"wp-caption-text\">Figure 9.11: A map of the world during the early Eocene showing one hypothesis for the direction of dispersal of the omomyoid Teilhardina. The map depicts primates hopping from continent to continent (East to West) via the forest corridors at high latitudes. Credit: <a href=\"https:\/\/www.pnas.org\/content\/103\/30\/11223\">Paleogeographic map showing hypothetical migration routes of Teilhardina (Figure 1)<\/a> by Thierry Smith, Kenneth D. Rose, and Philip D. Gingerich. 2006. <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">Proceedings of the National Academy of Sciences of the United States of America <\/a>103 (30): 11223\u201311227. Copyright (2006) National Academy of Sciences. Image <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">is used for non-commercial and educational purposes as outlined by PNAS.<\/a><\/figcaption><\/figure>\n<h2 class=\"import-Normal\">The Emergence of Modern Primate Groups<\/h2>\n<h3 class=\"import-Normal\"><strong>Origins of Crown Strepsirrhines<\/strong><\/h3>\n<p class=\"import-Normal\">Until the turn of this century, very little was known about the origins of the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1698\">crown<\/a><\/strong> (living) strepsirrhines. The Quaternary record of Madagascar contains many amazing forms of lemurs, including giant sloth-like lemurs, lemurs with perhaps monkey-like habits, lemurs with koala-like habits, and even a giant aye-aye (Godfrey and Jungers 2002). However, in Madagascar, early Tertiary continental sediments are lacking, and there is no record of lemur fossils before the Pleistocene.<\/p>\n<p class=\"import-Normal\">The fossil record of galagos is slightly more informative. Namely, there are Miocene African fossils that are very likely progenitors of lorisids (Simpson 1967). However, these are much like modern galagos and do not reveal anything about the relationship between crown strepsirrhines and Eocene fossil primates (but see below regarding <em>Propotto<\/em>). A similar situation exists for lorises in Asia: there are Miocene representatives, but these are substantially like modern lorises. The discovery of the first definite <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1699\"><strong>toothcomb<\/strong><\/a> canine (a hallmark of stresirrhines) in 2003 provided the \u201csmoking gun\u201d for the origin of crown strepsirrhines (Seiffert Et al. 2003). Recently, several other African primates have been recognized as having strepsirrhine affinities (Marivaux Et al. 2013; Seiffert 2012). The enigmatic Fayum primate <em>Plesiopithecus<\/em> is known from a skull that has been compared to aye-ayes and to lorises (Godinot 2006; Simons and Rasmussen 1994a).<\/p>\n<p class=\"import-Normal\">The now-recognized diversity of stem strepsirrhines from the Eocene and Oligocene of Afro-Arabia is strong evidence to suggest that strepsirrhines originated in that geographic area. This implies that lorises dispersed to Asia subsequent to an African origin. It is unknown what the first strepsirrhines in Madagascar were like. However, it seems likely that the lemuriform-lorisiform split occurred in continental Africa, followed by dispersal of lemuriform stock to Madagascar. Recent evidence suggests that <em>Propotto<\/em>, a Miocene primate from Kenya originally described as a potto antecedent, actually forms a clade with <em>Plesiopithecus <\/em>and the aye-aye; this might suggest that strepsirrhines dispersed to Madagascar from continental Africa more than once (Gunnell Et al. 2018).<\/p>\n<h3 class=\"import-Normal\"><strong>The Fossil Record of Tarsiers<\/strong><\/h3>\n<p class=\"import-Normal\">Tarsiers are so unusual that they fuel major debates about primate taxonomy. Tarsiers today are moderately diverse but geographically limited and not very different in their ecological habits\u2014especially considering that the split between them and their nearest living relative probably occurred over 50 million years ago. If omomyoids are excluded, then the fossil record of tarsiers is very limited. Two fossil species from the Miocene of Thailand have been placed in the genus <em>Tarsius<\/em>, as has an Eocene fossil from China (Beard Et al. 1994). These, and <em>Xanthorhysis<\/em> from the Eocene of China, are all very tarsier-like. In fact, it is striking that <em>Tarsius eocaenus<\/em> from China was already so tarsier-like as early as the Eocene. This suggests that tarsiers achieved their current morphology very early in their evolution and have remained more or less the same while other primates changed dramatically. Two additional genera, <em>Afrotarsius<\/em> from the Oligocene of Egypt and Libya and <em>Afrasia<\/em> from the Eocene of Myanmar, have also been implicated in tarsier origins, though the relationship between them and tarsiers is unclear (Chaimanee Et al. 2012). More recently, a partial skeleton of a small Eocene primate from China, <em>Archicebus achilles<\/em> (dated to approximately 55.8 million to 54.8 million years ago), was described as the most basal tarsiiform (Ni Et al. 2013). This primate is reconstructed as a diurnal insectivore and an arboreal quadruped that did some leaping\u2014but not to the specialized degree seen in living tarsiers. The anatomy of the eye in living tarsiers suggests that their lineage passed through a diurnal stage, so <em>Archicebus<\/em> (and diurnal omomyoids) might represent such a stage.<\/p>\n<h3 class=\"import-Normal\"><strong>Climate Change and the Paleogeography of Modern Primate Origins<\/strong><\/h3>\n<p class=\"import-Normal\">Changing global climate has had profound effects on primate dispersal patterns and ecological habits over evolutionary time. Primates today are strongly tied to patches of trees and particular plant parts such as fruits, seeds, and immature leaves. It is no surprise, then, that the distribution of primates mirrors the distribution of forests. Today, primates are most diverse in the tropics, especially in tropical rainforests. Global temperature trends across the Tertiary have affected primate ranges. Following the Cretaceous-Tertiary extinction event, cooler temperatures and greater seasonality characterized the Paleocene. In the Eocene, temperatures (and probably rainfall) increased globally and rainforests likely extended to very high latitudes. During this time, euprimates became diverse. With cooling and increased aridity at the end of the Eocene, many primate extinctions occurred in the northern continents and the surviving primates were confined to lower latitudes in South America, Afro-Arabia, Asia, and southern Europe. Among these survivors are the progenitors of the living groups of primates: lemurs and lorises, tarsiers, <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1700\"><strong>platyrrhines<\/strong><\/a> (monkeys of the Americas), and catarrhines (monkeys and apes of Africa and Asia) (Figure 9.12).<\/p>\n<figure id=\"attachment_277-14\" aria-describedby=\"caption-attachment-277-14\" style=\"width: 539px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-264\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image2-5-e1691791570984.png\" alt=\"Map of world with gray continents.\" width=\"539\" height=\"306\" \/><figcaption id=\"caption-attachment-277-14\" class=\"wp-caption-text\">Figure 9.12: Map of key localities of early anthropoids on land that becomes Africa and southern Asia. Credit: <a class=\"rId56\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Oligocene Map with Key Early Anthropoid Localities (Figure 8.10)<\/a> original to <a class=\"rId57\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Elyssa Ebding at <a class=\"rId58\" href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\">GeoPlace, California State University, Chico<\/a> is under a <a class=\"rId59\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Localities based on Fleagle 2013, 265.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Competing Hypotheses for the Origin of Anthropoids<\/strong><\/h3>\n<p class=\"import-Normal\">There is considerable debate among paleoanthropologists as to the geographic origins of anthropoids. In addition, there is debate regarding the source group for anthropoids. Three different hypotheses have been articulated in the literature. These are the adapoid origin hypothesis, the omomyoid origin hypothesis, and the tarsier origin hypothesis (Figure 9.13).<\/p>\n<figure style=\"width: 419px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image24-1-1.jpg\" alt=\"Diagrams show three relationships among primate groups.\" width=\"419\" height=\"742\" \/><figcaption class=\"wp-caption-text\">Figure 9.13: Competing models of anthropoid origins. Branch lengths are not to scale. The omomyoid origin model and tarsier origin model do not make specific reference to the evolutionary position of strepsirrhines; however, they were included here for completeness. <a href=\"https:\/\/docs.google.com\/document\/d\/1VUDKMBJYS_jNONjLxT04jQN0_z9Ua50BRN6auGSHUuU\/edit\">A full text description of this image is available<\/a>. Credit: <a class=\"rId61\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Competing Trees for Anthropoid Origins (Figure 8.11)<\/a> original to <a class=\"rId62\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Jonathan M. G. Perry is under a <a class=\"rId63\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Adapoid Origin Hypothesis<\/em><\/h4>\n<p class=\"import-Normal\">Resemblances between some adapoids and some extant anthropoids include fusion of the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1702\"><strong>mandibular symphysis<\/strong><\/a>, overall robusticity of the chewing system, overall large body size, features that signal a diurnal lifestyle (like relatively small eye sockets), and ankle bone morphology. Another feature in common is canine sexual dimorphism, which is present in some species of adapoids (probably) and in several species of anthropoids.<\/p>\n<p class=\"import-Normal\">These features led some paleoanthropologists in the last half of the 20th century to suggest that anthropoids came from adapoid stock (Gingerich 1980; Simons and Rasmussen 1994b). One of the earliest supporters of the link between adapoids and anthropoids was Hans Georg Stehlin, who described much of the best material of adapoids and compared these Eocene primates to South American monkeys (Stehlin 1912). In more recent times, the adapoid origin hypothesis was reinforced by resemblances between these European adapoids (especially <em>Adapis <\/em>and <em>Leptadapis<\/em>) and some early anthropoids from the Fayum Basin (e.g., <em>Aegyptopithecus<\/em>, see below; Figure 9.14).<\/p>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 473.25pt\">\n<caption>Figure 9.14: Families of early anthropoids with example genera and traits: a table. Credit: Early anthropoids table original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Jonathan M. G. Perry and Stephanie L. Canington is under a <a class=\"rId64\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Content derived from Fleagle 2013.<\/caption>\n<thead>\n<tr style=\"height: 25pt\">\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Family<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Genera<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Morphology<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Location<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Age<\/strong><sup><strong>1<\/strong><\/sup><\/p>\n<p>&nbsp;<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table3-R\" style=\"height: 18pt\">\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Propliopithecidae<sup>2<\/sup><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Aegyptopithecus<\/em><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Large size. Cranial sexual dimorphism, large canines. Robust jaws and rounded molars. Partially ossified ear tube (in some). Robust skeleton; quadruped.<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Africa<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Late Eocene to Early Oligocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 16pt\">\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Parapithecidae<sup>3<\/sup><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Apidium<\/em><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Medium size. Retention of three premolars per quadrant. Rounded molars and premolars with large central cusps. Adaptations for leaping in the lower limb.<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Africa<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Late Eocene to Late Oligocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 16pt\">\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Proteopithecidae<sup>4<\/sup><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Proteopithecus<\/em><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Small size. Retention of three premolars per quadrant. Arboreal quadrupeds that ate fruit.<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Africa<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Late Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 16pt\">\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Oligopithecidae<sup>5<\/sup><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Catopithecus<\/em><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Small size. Skull has postorbital septum and unfused mandible. Deep jaws. Diet of fruits. Generalized quadruped.<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Africa<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Late Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 16pt\">\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Eosimiidae<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Eosimias<\/em><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Deep jaw with vertical unfused symphysis. Pointed incisors and canines. Crowded premolars.<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Asia<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Middle Eocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 16pt\">\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Amphipithecidae<sup>6<\/sup><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\"><em>Pondaungia<\/em><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Deep jaws. Molars generally rounded with wide basins.<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Asia<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt 0pt 5.4pt;border: solid #000000 0.5pt\">\n<p class=\"import-Normal\">Middle Eocene to Early Oligocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 1pt\">\n<td class=\"Table3-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\" colspan=\"4\">\n<p class=\"import-Normal\"><sup>1<\/sup> Derived from Fleagle 2013.<\/p>\n<p class=\"import-Normal\"><sup>2<\/sup> See Gebo and Simons 1987 and Simons Et al. 2007.<\/p>\n<p class=\"import-Normal\"><sup>3<\/sup> See Feagle and Simons 1995 and Simons 2001.<\/p>\n<p class=\"import-Normal\"><sup>4<\/sup> See Simons and Seiffert 1999.<\/p>\n<p class=\"import-Normal\"><sup>5<\/sup> See Simons and Rasmussen 1996.<\/p>\n<p class=\"import-Normal\"><sup>6<\/sup> See Kay Et al. 2004.<\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"border-top: solid #000000 0.5pt;border-right: none #000000 0pt;border-bottom: none #000000 0pt;border-left: none #000000 0pt;padding: 0pt 5.4pt 0pt 5.4pt\">\n<p class=\"import-Normal\">\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<p class=\"import-Normal\">Unfortunately for the adapoid hypothesis, most of the shared features listed above probably emerged independently in the two groups as adaptations to a diet of hard and\/or tough foods. For example, fusion of the mandibular symphysis likely evolved as a means to strengthen the jaw against forces that would pull the two halves away from each other, in the context of active chewing muscles on both sides of the head generating great bite forces. This context would also favour the development of robust jaws, large chewing muscles, shorter faces, and some other features shared by some adapoids and some anthropoids.<\/p>\n<p class=\"import-Normal\">As older and more plesiomorphic anthropoids were found in the Fayum Basin, it became clear that the earliest anthropoids from Africa did not possess these features of jaw robusticity (Seiffert Et al. 2009). Furthermore, many adapoids never evolved these features. Fusion of the mandibular symphysis in adapoids is actually quite different from that in anthropoids and probably occurred during juvenile development in the former (Beecher 1983; Ravosa 1996). Eventually, the adapoid origin hypothesis fell out of favour among most paleoanthropologists, although the description of <em>Darwinius<\/em> is a recent revival of that idea (Franzen Et al. 2009; but see Seiffert Et al. 2009, Williams Et al. 2010b).<\/p>\n<h4 class=\"import-Normal\"><em>Omomyoid Origin Hypothesis<\/em><\/h4>\n<p class=\"import-Normal\">Similarities in cranial and hindlimb morphology between some omomyoids and extant tarsiers have led to the suggestion that tarsiers arose from some kind of omomyoid. In particular, <em>Necrolemur<\/em> has many features in common with tarsiers, as does the North American <em>Shoshonius<\/em>, which is known from a few beautifully preserved (although distorted) crania. Tarsiers and <em>Shoshonius <\/em>share exclusively some features of the base of the cranium; however, <em>Shoshonius<\/em> does not have any sign of postorbital closure, and it lacks the bony ear tube of tarsiers. Nevertheless, some of the resemblances between some omomyoids and tarsiers suggest that tarsiers might have originated from within the Omomyoidea (Beard 2002; Beard and MacPhee 1994). In this scenario, although living tarsiers and living anthropoids might be sister taxa, they might have evolved from different omomyoids, possibly separated from each other by more than 50 million years of evolution, or from anthropoids evolved from some non-omomyoid fossil group. The arguments against the omomyoid origin hypothesis are essentially the arguments <em>for<\/em> the tarsier origin hypothesis (see below). Namely, tarsiers and anthropoids share many features (especially of the soft tissues) that must have been retained for many millions of years or must have evolved convergently in the two groups. Furthermore, a key hard-tissue feature shared between the two extant groups, the postorbital septum, was not present in any omomyoid. Therefore, that feature must have arisen convergently in the two extant groups or must have been lost in omomyoids. Neither scenario is very appealing, although recent arguments for <strong>convergent evolution<\/strong> of the postorbital septum in tarsiers and anthropoids have arisen from embryology and histology of the structure (DeLeon Et al. 2016).<\/p>\n<h4 class=\"import-Normal\"><em>Tarsier Origin Hypothesis<\/em><\/h4>\n<p class=\"import-Normal\">Several paleoanthropologists have suggested that there is a relationship between tarsiers and anthropoids to the exclusion of omomyoids and adapoids (e.g., Cartmill and Kay 1978; Ross 2000; Williams and Kay 1995). Tarsiers and anthropoids today share several traits, including many soft-tissue features related to the olfactory system (e.g., the loss of a hairless external nose and loss of the median cleft running from the nose to the mouth, as possessed by strepsirrhines), and aspects of the visual system (e.g., the loss of a reflective layer at the back of the eye, similarities in carotid circulation to the brain, and mode of placentation). Unfortunately, none of these can be assessed directly in fossils. Some bony similarities between tarsiers and anthropoids include an extra air-filled chamber below the middle ear cavity, reduced bones within the nasal cavity, and substantial postorbital closure; these can be assessed in fossils, but the distribution of these traits in omomyoids does not yield clear answers. Furthermore, several similarities between tarsiers and anthropoids are probably due to similarities in sensory systems, which might have evolved in parallel for ecological reasons. Although early attempts to resolve the crown primates with molecular data were sometimes equivocal or in disagreement with one another, more recent analyses (including those of short interspersed elements) suggest that tarsiers and anthropoids are sister groups to the exclusion of lemurs and lorises (Williams Et al. 2010a). However, this does not address omomyoids, all of which are far too ancient for DNA extraction.<\/p>\n<p class=\"import-Normal\">The above three hypotheses are not the only possibilities for anthropoid origins. It may be that anthropoids are neither the closest sister group of tarsiers, nor evolved from adapoids or omomyoids. In recent years, two new groups of Eocene Asian primates have been implicated in the origin of anthropoids: the eosimiids and the amphipithecids. It is possible that one or the other of these two groups gave rise to anthropoids. Regardless of the true configuration of the tree for crown primates, the three major extant groups probably diverged from each other quite long ago (Seiffert Et al. 2004).<\/p>\n<h3 class=\"import-Normal\"><strong>Early Anthropoid Fossils in Africa<\/strong><\/h3>\n<figure style=\"width: 526px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image7-2.jpg\" alt=\"People digging in a sandy desert.\" width=\"526\" height=\"352\" \/><figcaption class=\"wp-caption-text\">Figure 9.15: Egyptian workers sweeping Quarry I in the Fayum Basin (2004). This technique, called wind harvesting, removes the desert crust and permits wind to blow out fine sediment and reveal fossils. Credit: <a class=\"rId66\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Egyptian workers sweeping Quarry I in the Fayum Basin (2004, Figure 8.12)<\/a> by Jonathan M. G. Perry is under a <a class=\"rId67\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<figure style=\"width: 280px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image14-2.jpg\" alt=\"A person using a tool to expose bone in sand.\" width=\"280\" height=\"423\" \/><figcaption class=\"wp-caption-text\">Figure 9.16: Elwyn Laverne Simons excavating Aegyptopithecus in the Fayum Basin. Credit: <a class=\"rId69\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Elwyn Laverne Simons in the Fayum Basin (Figure 8.13)<\/a> used by permission of the <a class=\"rId70\" href=\"https:\/\/lemur.duke.edu\/\">Duke Lemur Center,<\/a> Division of Fossil Primates, is under a <a class=\"rId71\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">The classic localities yielding the greatest wealth of early anthropoid fossils are those from the Fayum Basin in Egypt (Simons 2008; Figure 9.15). The Fayum is a veritable oasis of fossil primates in an otherwise spotty early Tertiary African record. Since the 1960s, teams led by E. L. Simons have discovered several new species of early anthropoids, some of which are known from many parts of the skeleton and several individuals (Figure 9.16).<\/p>\n<p class=\"import-Normal\">The Fayum Jebel Qatrani Formation and Birket Qarun Formation between them have yielded a remarkable array of terrestrial, arboreal, and aquatic mammals. These include ungulates, bats, sea cows, elephants, hyraces, rodents, whales, and primates. Also, many other vertebrates, like water birds, were present. The area at the time of deposition (Late Eocene through Early Oligocene) was probably very wet, with slow-moving rivers, standing water, swampy conditions, and lots of trees (see Bown and Kraus 1988). In short, it was an excellent place for primates.<\/p>\n<h4 class=\"import-Normal\"><em>General Morphology of Anthropoids<\/em><\/h4>\n<p class=\"import-Normal\">The anthropoids known from the Fayum (and their close relatives from elsewhere in East Africa and Afro-Arabia) bear many of the anatomical hallmarks of extant anthropoids; however, there are plesiomorphic forms in several families that lack one or more anthropoid traits. All Fayum anthropoids known from skulls possess postorbital closure, most had fused mandibular symphyses, and most had ring-like <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1704\"><strong>ectotympanic<\/strong> <\/a> bones. Tooth formulae were generally either 2.1.3.3 or 2.1.2.3. Fayum anthropoids ranged in size from the very small <em>Qatrania<\/em> and <em>Biretia <\/em>(less than 500 g) to the much-larger <em>Aegyptopithecus<\/em> (approximately 7 kg; 15 lbs.). Fruit was probably the main component of the diet for most or all of the anthropoids, with some of them supplementing with leaves (Kay and Simons 1980; Kirk and Simons 2001; Teaford Et al. 1996). Most Fayum anthropoids were probably diurnal above-branch quadrupeds. Some of them (e.g., <em>Apidium<\/em>; see Figure 9.14) were probably very good leapers (Gebo and Simons 1987), but none show specializations for gibbon-style suspensory locomotion. Some of the Fayum anthropoids are known from hundreds of individuals, permitting the assessment of individual variation, sexual dimorphism, and in some cases growth and development. The description that follows provides greater detail for the two best known Fayum anthropoid families, the Propliopithecidae and the Parapithecidae; the additional families are summarized briefly.<\/p>\n<h4 class=\"import-Normal\"><em>Fayum Anthropoid Families<\/em><\/h4>\n<p class=\"import-Normal\">The Propliopithecidae (see Figure 9.14) include the largest anthropoids from the fauna, and they are known from several crania and some postcranial elements. They have been suggested to be stem catarrhines, although perhaps near the split between catarrhines and platyrrhines. The best known propliopithecid is <em>Aegyptopithecus<\/em>, known from many teeth, crania, and postcranial elements (Figure 9.17) .<\/p>\n<figure style=\"width: 431px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-2-1.jpg\" alt=\"Two animal skull side views.\" width=\"431\" height=\"281\" \/><figcaption class=\"wp-caption-text\">Figure 9.17: Female (left) and male (right) skull material for Aegyptopithecus zeuxis. The mandibles are not associated with the crania. Credit: <a href=\"https:\/\/www.pnas.org\/doi\/full\/10.1073\/pnas.0703129104#supplementary-materials\">Female and male cranium of A. zeuxi (03129Fig5, Supporting Information)<\/a> by Elwyn L. Simons, Erik R. Seiffert, Timothy M. Ryan, and Yousry Attia. 2007. <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">Proceedings of the National Academy of Sciences of the United States of America<\/a> 104 (21): 8731\u20138736. Copyright (2007) National Academy of Sciences. Image <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">is used for non-commercial and educational purposes as outlined by PNAS.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\">Parapithecidae are an extremely abundant and unusual family of anthropoids from the Fayum. The parapithecid <em>Apidium<\/em> is known from many jaws with teeth, crushed and distorted crania, and several skeletal elements. <em>Parapithecus<\/em> is known from cranial material including a beautiful, undistorted cranium. This genus shows extreme reduction of the incisors, including complete absence of the lower incisors in <em>P. grangeri <\/em>(Simons 2001). This trait is unique among primates. Parapithecids were once thought to be the ancestral stock of platyrrhines; however, their platyrrhine-like features are probably ancestral retentions, so the most conservative approach is to consider them stem anthropoids.<\/p>\n<p class=\"import-Normal\">The Proteopithecidae were small frugivores that probably mainly walked along horizontal branches on all fours. They are considered stem anthropoids. The best known genus, <em>Proteopithecus<\/em>, is represented by dentitions, crania, and post-cranial elements.<\/p>\n<p class=\"import-Normal\">The Oligopithecidae share a mixture of traits that makes them difficult to classify more specifically within anthropoids. The best known member, <em>Catopithecus<\/em>, is known from crania that demonstrate a postorbital septum and from mandibles that lack symphyseal fusion. They share the catarrhine tooth formula of 2.1.2.3 and have a canine honing complex that involves the anterior lower premolar. The postcranial elements known for the group suggest generalized arboreal quadrupedalism. The best known member, <em>Catopithecus<\/em>, is known from crania that demonstrate a postorbital septum and from mandibles that lack symphyseal fusion (Simons and Rasmussen 1996). The jaws are deep, with broad muscle attachment areas and crested teeth. <em>Catopithecus<\/em> was probably a little less than a kilogram in weight.<\/p>\n<p class=\"import-Normal\">Other genera of putative anthropoids from the Fayum include the very poorly known <em>Arsinoea<\/em>, the contentious <em>Afrotarsius<\/em>, and the enigmatic <em>Nosmips<\/em>. The last of these possesses traits of several major primate <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1705\">clades<\/a><\/strong> and defies classification (Seiffert Et al. 2010).<\/p>\n<h3 class=\"import-Normal\"><strong>Early Anthropoid Fossils in Asia<br style=\"clear: both\" \/><\/strong><\/h3>\n<p class=\"import-Normal\">For the last half of the 1900s, researchers believed that Africa was the unquestioned homeland of early anthropoids (see Fleagle and Kay 1994). However, two very different groups of primates from Asia soon began to change that. One was an entirely new discovery (Eosimiidae), and the other was a poorly known group discovered decades prior (Amphipithecidae). Soon, attention on anthropoid origins began to shift eastward (see Ross and Kay 2004; Simons 2004). If anthropoids arose in Asia instead of Africa, then this implies that the African early anthropoids either emigrated from Asia or evolved their anthropoid traits in parallel with living anthropoids.<\/p>\n<h4 class=\"import-Normal\"><em>Eosimiids<\/em><\/h4>\n<p class=\"import-Normal\">First described in the 1990s, the eosimiids are best represented by <em>Eosimias <\/em>(see Figure 9.14; Figure 9.18). This tiny \u201cdawn monkey\u201d is known from relatively complete jaws with teeth, a few small fragments of the face, and some postcranial elements (Beard Et al. 1994; Beard Et al. 1996; Gebo Et al. 2000). <em>Eosimias<\/em> (along with the other less-well-known genera in its family) bears some resemblance to tarsiers as well as anthropoids. Unfortunately, no good crania are known for this family, and the anatomy of, for example, the posterior orbital margin could be very revealing as to higher-level relationships.<\/p>\n<figure style=\"width: 550px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image16-1-1.jpg\" alt=\"Red-colored lower jaw of an animal.\" width=\"550\" height=\"232\" \/><figcaption class=\"wp-caption-text\">Figure 9.18: Cast of the right half of the mandible of Eosimias centennicus, type specimen. The white scale bar is 1 cm long. Credit: <a class=\"rId74\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Cast of the right half of the mandible of <\/a><a class=\"rId75\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\"><em>Eosimias centennicus <\/em><\/a><a class=\"rId76\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">(Figure 8.15),<\/a> type specimen, from K. D. Rose cast collection, photo by Jonathan M. G. Perry is under a <a class=\"rId77\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Amphipithecids<\/em><\/h4>\n<p class=\"import-Normal\">Amphipithecids are small- to medium-size primates (up to 10 kg; 22 lbs.). Most are from the Eocene Pondaung Formation in Myanmar (Early\u2013Middle Eocene), but one genus is known from Thailand. Some dental similarities with anthropoids were noted early on, such as deep jaws and wide basins that separate low molar cusps. The best known genera were <em>Pondaungia<\/em> and <em>Amphipithecus <\/em>(Ciochon and Gunnell 2002; see Figure 9.14). Another amphipithecid, <em>Siamopithecus<\/em> from Thailand, has very rounded molars and was probably a seed-eater (Figure 9.19). In addition to teeth and jaws, some cranial fragments, ankle material, and ends of post-cranial bones have been found for <em>Pondaungia<\/em>. There are important resemblances between these bones of <em>Pondaungia<\/em> and those of adapoids, suggesting adapoid affinities for the amphipithecidae. This would imply that the resemblances with anthropoids in the teeth are convergent, based on similarities in diet (see Ciochon and Gunnell 2002). Unfortunately, the association between post-cranial bones and teeth is not definite. With other primates in these faunas (including eosimiids), one cannot be certain that the post-cranial bones belong with the teeth. Some researchers suggest that some bones belong to a sivaladapid (or asiadapid) and others to an early anthropoid (Beard Et al. 2007; Marivaux Et al. 2003). Additional well-associated material of amphipithecids would help to clear up this uncertainty.<\/p>\n<figure style=\"width: 505px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image15-2.jpg\" alt=\"Four casts of jawbone fragments with teeth.\" width=\"505\" height=\"368\" \/><figcaption class=\"wp-caption-text\">Figure 9.19: Casts of representative amphipithecid material. A. Pondaungia cotteri right lower jaw fragment with m2 and m3. B. Siamopithecus eocaenus right upper jaw fragment with p4-m3. C. S. eocaenus right lower jaw fragment with partial m1, m2, and m3 in lateral view. D. Same as in C but occlusal view. White scale bars are 1 cm long. Credit: <a class=\"rId79\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Casts of representative amphipithecid material (Figure 8.16)l<\/a> from K. D. Rose cast collection, photo by Jonathan M. G. Perry is under a <a class=\"rId80\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Platyrrhine Dispersal to South America<\/strong><\/h3>\n<p class=\"import-Normal\">Today there is an impressive diversity of primates in South and Central America. These are considered to be part of a single clade, the Platyrrhini. Primates colonized South America sometime in the Eocene from an African source. In the first half of the 20th century, the source of platyrrhines was a matter of major debate among paleontologists, with some favouring a North American origin (e.g., Simpson 1940).<\/p>\n<p class=\"import-Normal\">Part of the reason for this debate is that South America was an island in the Eocene. Primates needed to cross open ocean to get there from either North America or Africa, although the distance from the former was shorter. Morphology yields clues to platyrrhine origins. The first known primates in South America have more in common morphologically with African primates than with North American ones. At the time, anthropoids were popping up in North Africa, whereas the only euprimates in North America were adapoids and omomyoids. Despite lacking a bony ear tube, early platyrrhines shared a great deal with other anthropoids, including full postorbital closure and fusion of the mandibular symphysis.<\/p>\n<p class=\"import-Normal\">The means by which a population of small North African primates managed to disperse across the Atlantic and survive to colonize South America remains a mystery. The most plausible scenario is one of rafting. That is, primates must have been trapped on vegetation that was blown out to sea by a storm. The vegetation then became a sort of life raft, which eventually landed ashore, dumping its passengers in South America. Rodents probably arrived in South America in the same way (Antoine Et al. 2012).<\/p>\n<p class=\"import-Normal\">Once ashore, platyrrhines must have crossed South America fairly rapidly because the earliest-known primates from that continent are from Peru (Bond Et al. 2015). Soon after that, platyrrhines were in Bolivia, namely <em>Branisella<\/em>. By the Miocene, platyrrhines were living in extreme southern Argentina and were exploiting a variety of feeding niches. The Early Miocene platyrrhines were all somewhat plesiomorphic in their morphology, but some features that likely arose by ecological convergence suggest (to some) relationships with extant platyrrhine families. This has led to a lively debate about the pattern of primate evolution in South America (Kay 2015; Kay &amp; Fleagle 2010; Rosenberger 2010). By the Middle Miocene, clear representatives of modern families were present in a diverse fauna from La Venta, Colombia (Wheeler 2010). The Plio-Pleistocene saw the emergence of giant platyrrhines as well as several taxa of platyrrhines living on Caribbean islands (Cooke Et al. 2016).<\/p>\n<p class=\"import-Normal\">The story of platyrrhines seems to be one of amazing sweepstakes dispersal, followed by rapid diversification and widespread geographic colonization of much of South America. After that, dramatic extinctions resulted in the current, much-smaller geographic distribution of platyrrhines. These extinctions were probably caused by changing climates, leading to the contraction of forests. Platyrrhines dispersed to the Caribbean and to Central America, with subsequent extinctions in those regions that might have been related to interactions with humans. Unlike anthropoids of Africa and Asia, platyrrhines do not seem to have evolved any primarily terrestrial forms and so have always been highly dependent on forests.<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Jonathan Perry and Primates of the Extreme South<\/h2>\n<p class=\"import-Normal\">Many primates are very vulnerable to ecological disturbance because they are heavily dependent on fruit to eat and trees to live in. This is one reason why so many primates are endangered today and why many of them went extinct due to climatic and vegetational changes in the past. I (Jonathan Perry) have conducted paleontological research focusing on primates that lived on the edge of their geographic distribution. This research has taken me to extreme environments in the Americas: southern Patagonia, the Canadian prairies, western Wyoming, and the badlands of eastern Oregon.<\/p>\n<p class=\"import-Normal\">Santa Cruz Province in Argentina is as far south as primates have ever lived. The Santa Cruz fauna of the Miocene has yielded a moderate diversity of platyrrhines, each with slightly different dietary adaptations. These include <em>Homunculus<\/em>, first described by Florentino Ameghino in 1891 (Figure 9.20). Recent fieldwork by my colleagues and I in Argentina has revealed several skulls of <em>Homunculus <\/em>as well as many parts of the skeleton (Kay Et al. 2012). The emerging profile of this extinct primate is one of a dedicated arboreal quadruped that fed on fruits and leaves. Many of the foods eaten by <em>Homunculus<\/em> must have been very tough and were probably covered and impregnated with grit; we suspect this because the cheek teeth are very worn down, even in young individuals, and because the molar tooth roots were very large, presumably to resist strong bite forces (Perry Et al. 2010, 2014).<\/p>\n<figure style=\"width: 497px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image9-2.jpg\" alt=\"An animal skull, a partial skull, and a fossil jaw with teeth.\" width=\"497\" height=\"634\" \/><figcaption class=\"wp-caption-text\">Figure 9.20: Representative specimens of Homunculus patagonicus. A. Adult cranium in lateral view. B. Adult cranium surface reconstructed from microCT scans, with the teeth segmented out. C. Juvenile cranium. White scale bars are 1cm long. Credit: <a class=\"rId82\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Representative specimens of <\/a><a class=\"rId83\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\"><em>Homunculus patagonicus <\/em><\/a><a class=\"rId84\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">(Figure 8.17)<\/a> photo by Jonathan M. G. Perry is under a <a class=\"rId85\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">I began working in Argentina while a graduate student at Duke University. I participated as a field assistant in a team led by my Ph.D. advisor, Richard F. Kay, and Argentine colleagues Sergio F. Vizca\u00edno and M. Susana Bargo. Most of the localities examined belong to a suite of beach sites known since the 1800s and visited by many field parties from various museums in the early 1900s. Since 2003, our international team of paleontologists from the U.S. and Argentina has visited these localities every single year (Figure 9.21). Over time, new fossils and new students have led to new projects and new approaches, including the use of microcomputed tomography (microCT) to visualize and analyse internal structures of the skeleton.<\/p>\n<figure style=\"width: 491px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image23.jpg\" alt=\"Sandy rocky coastline. People digging on a grassy hillside.\" width=\"491\" height=\"561\" \/><figcaption class=\"wp-caption-text\">Figure 9.21: Field localities in Argentina and Canada. A. Ca\u00f1adon Palos locality, coastal Santa Cruz Province, Argentina. B. Swift Current Creek locality, southwest Saskatchewan, Canada. Credits: A. <a class=\"rId87\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Ca\u00f1adon Palos Field Locality in Argentina<\/a> by Jonathan M. G. Perry is under a <a class=\"rId88\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. B. <a class=\"rId89\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Swift Current Creek locality, Saskatchewan, Canada<\/a> by Jonathan M. G. Perry is under a <a class=\"rId90\" 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\">Planet of Apes<\/h2>\n<h3 class=\"import-Normal\"><strong>Geologic Activity and Climate Change in the Miocene<\/strong><\/h3>\n<p class=\"import-Normal\">The Miocene Epoch was a time of mammalian diversification and extinction, global climate change, and ecological turnover. In the Miocene, there was an initial warming trend across the globe with the expansion of subtropical forests, followed by widespread cooling and drying with the retreat of tropical forests and replacement with more open woodlands and eventually grasslands. It was also a time of major geologic activity. On one side of the globe, South America experienced the rise of the Andes Mountains. On the other side, the Indian subcontinent collided with mainland Asia, resulting in the rise of the Himalayan Mountains. In Africa, volcanic activity promoted the development of the East African Rift System. Critical to the story of ape evolution was the exposure of an intercontinental landbridge between East Africa and Eurasia, permitting a true planet of apes (Figure 9.22).<\/p>\n<figure id=\"attachment_277-15\" aria-describedby=\"caption-attachment-277-15\" style=\"width: 580px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-273\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image21-3-e1691792198797.png\" alt=\"Map of world with gray continents.\" width=\"580\" height=\"335\" \/><figcaption id=\"caption-attachment-277-15\" class=\"wp-caption-text\">Figure 9.22: Map of the world in the Miocene, highlighting fossil ape localities across Africa, southern Europe, and southern Asia. Credit: <a class=\"rId92\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-10\/\">Miocene Map with Fossil Ape Localities (Figure 8.19)<\/a> original to <a class=\"rId93\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Elyssa Ebding at <a class=\"rId94\" href=\"https:\/\/www.csuchico.edu\/geop\/geoplace\/index.shtml\">GeoPlace, California State University, Chico<\/a> is under a <a class=\"rId95\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Localities based on Fleagle 2013, 311.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Geographic Distribution: Africa, Asia, Europe<\/strong><\/h3>\n<p class=\"import-Normal\">The world of the Miocene had tremendous ape diversity compared to today. The earliest records of fossil apes are from Early Miocene deposits in Africa. However, something dramatic happened around 16 million years ago. With the closure of the ancient Tethys Sea, the subsequent exposure of the <em>Gomphotherium<\/em> Landbridge, and a period of global warming, the Middle\u2013Late Miocene saw waves of emigration of mammals (including primates) out of Africa and into Eurasia, with evidence of later African re-entry for some (Harrison 2010). Some of the mammals that dispersed from Africa to Eurasia and back were apes. Though most of these early apes left no modern descendants, some of them gave rise to the ancestors of modern apes\u2014including <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_800\">hominins<\/a><\/strong> (Figure 9.23).<\/p>\n<figure style=\"width: 560px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image20-1.jpg\" alt=\"Miocene apes set against a geologic time scale.\" width=\"560\" height=\"796\" \/><figcaption class=\"wp-caption-text\">Figure 9.23: Representative Miocene apes set against a geologic time scale. Credit: <a href=\"https:\/\/www.pnas.org\/content\/108\/14\/5554\">Range chart for Miocene hominoids of Western Eurasia (Figure 3)<\/a> by Isaac Casanovas-Vilar, David M. Alba, Miguel Garc\u00e9s, Josep M. Robles, and Salvador Moy\u00e0-Sol\u00e0. 2011. <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">Proceedings of the National Academy of Sciences of the United States of America<\/a> 108 (14): 5554-5559. Copyright (2011) National Academy of Sciences. Image <a href=\"https:\/\/www.pnas.org\/about\/rights-permissions\">is used for non-commercial and educational purposes as outlined by PNAS.<\/a><\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Where Are the Monkeys? Diversity in the Miocene<\/strong><\/h3>\n<p class=\"import-Normal\">Whereas the Oligocene deposits in the Fayum of Egypt have yielded the earliest-known catarrhine fossils, the Miocene demonstrates some diversification of Cercopithecoidea. However, compared to the numerous and diverse Miocene apes (see below), monkeys of the Miocene are very rare and restricted to a single extinct family, the Victoriapithecidae (Figure 9.24). This family contains the earliest definite cercopithecoids.<\/p>\n<figure style=\"width: 357px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image13-5.png\" alt=\"Front view of skull with pointed teeth.\" width=\"357\" height=\"322\" \/><figcaption class=\"wp-caption-text\">Figure 9.25: Skull of Victoriapithecus macinnesi. Credit: Victoriapithecus macinnesi skull photo taken at the Musee d'Histoire Naturelle, Paris by Ghedoghedo is under a CC BY-SA 3.0 License.<\/figcaption><\/figure>\n<p class=\"import-Normal\">These monkeys are found from northern and eastern Africa between 20 million and 12.5 million years ago (Miller Et al. 2009). The best known early African monkey is <em>Victoriapithecus <\/em>(Figure 9.25), a small-bodied (approximately 7 kg; 15 lbs.), small-brained monkey. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1708\">Bilophodonty<\/a><\/strong>, known to be a hallmark of molar teeth of modern cercopithecoid, was present to some extent in Victoriapithecids. <em>Victoriapithecus<\/em> has been reconstructed as being more frugivorous and perhaps spent more time on the ground (terrestrial locomotion) than in the trees (arboreal locomotion; Blue Et al. 2006). The two major groups of cercopithecoids today are cercopithecines and colobines. The earliest records demonstrating clear members of each of these two groups are at the end of the Miocene. Examples include the early colobine <em>Microcolobus<\/em> from Kenya and the early cercopithecine <em>Pliopapio<\/em> from Ethiopia.<\/p>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 473.25pt;height: 349px\">\n<caption>Figure 9.24: Some families of later anthropoids with example genera and traits: a table. Credit: Late anthropoids table original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Jonathan M. G. Perry and Stephanie L. Canington is under a <a class=\"rId100\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Content derived from Fleagle 2013.<\/caption>\n<thead>\n<tr style=\"height: 25pt\">\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 60px;width: 119.35px\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Family<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 60px;width: 103.417px\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Genera<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 60px;width: 191.65px\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Morphology<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 60px;width: 67.3667px\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Location<\/strong><\/p>\n<p>&nbsp;<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 60px;width: 73.2167px\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Age<\/strong><sup><strong>1<\/strong><\/sup><\/p>\n<p>&nbsp;<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table4-R\" style=\"height: 18pt\">\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 77px;width: 119.35px\">\n<p class=\"import-Normal\">Victoriapithecidae<sup>2<\/sup><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 77px;width: 103.417px\">\n<p class=\"import-Normal\"><em>Victoriapithecus<\/em><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 77px;width: 191.65px\">\n<p class=\"import-Normal\">Long, sloping face. Round, narrowly spaced orbits. Deep cheek bones. Well-developed sagittal crest.<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 77px;width: 67.3667px\">\n<p class=\"import-Normal\">Africa<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 77px;width: 73.2167px\">\n<p class=\"import-Normal\">Early to Middle Miocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 16pt\">\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 61px;width: 119.35px\">\n<p class=\"import-Normal\">Proconsulidae<sup>3<\/sup><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 61px;width: 103.417px\">\n<p class=\"import-Normal\"><em>Proconsul<\/em><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 61px;width: 191.65px\">\n<p class=\"import-Normal\">Short face. Generalized dentition. Arboreal quadruped. Probably tailless.<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 61px;width: 67.3667px\">\n<p class=\"import-Normal\">Africa and Arabia<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 61px;width: 73.2167px\">\n<p class=\"import-Normal\">Early to Middle Miocene<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 16pt\">\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 46px;width: 119.35px\">\n<p class=\"import-Normal\">Pongidae<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 46px;width: 103.417px\">\n<p class=\"import-Normal\"><em>Gigantopithecus<\/em><\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 46px;width: 191.65px\">\n<p class=\"import-Normal\">Largest primate ever. Deep jaws and low rounded molars.<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 46px;width: 67.3667px\">\n<p class=\"import-Normal\">Asia<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"vertical-align: middle;padding: 0pt 5.4pt;border: 0.5pt solid #000000;height: 46px;width: 73.2167px\">\n<p class=\"import-Normal\">Miocene to Present<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table4-R\" style=\"height: 1pt\">\n<td class=\"Table4-C\" style=\"border-color: #000000;border-style: solid none none;border-width: 0.5pt 0pt 0pt;padding: 0pt 5.4pt;height: 90px;width: 526.983px\" colspan=\"4\">\n<p class=\"import-Normal\"><sup>1<\/sup> Derived from Fleagle 2013.<\/p>\n<p class=\"import-Normal\"><sup>2<\/sup> See Benefit and McCrossin 1997 and Fleagle 2013.<\/p>\n<p class=\"import-Normal\"><sup>3<\/sup> See Begun 2007.<\/p>\n<\/td>\n<td class=\"Table4-C\" style=\"border-color: #000000;border-style: solid none none;border-width: 0.5pt 0pt 0pt;padding: 0pt 5.4pt;height: 90px;width: 73.2167px\">\n<p class=\"import-Normal\">\n<\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"height: 15px;width: 121.283px\"><\/td>\n<td style=\"height: 15px;width: 105.35px\"><\/td>\n<td style=\"height: 15px;width: 193.583px\"><\/td>\n<td style=\"height: 15px;width: 69.3px\"><\/td>\n<td style=\"height: 15px;width: 74.65px\"><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<h3><\/h3>\n<h3><\/h3>\n<h3><\/h3>\n<h3 class=\"import-Normal\"><strong>The Story of Us, the Apes<\/strong><\/h3>\n<h4 class=\"import-Normal\"><em>African Ape Diversity <\/em><\/h4>\n<p class=\"import-Normal\">The Early Miocene of Africa has yielded around 14 genera of early apes (Begun 2003). Many of these taxa have been reconstructed as frugivorous arboreal quadrupeds (Kay 1977). One of the best studied of these genera is the East African <em>Proconsul<\/em> (Family Proconsulidae; see Figure 9.24). Several species have been described, with body mass reconstructions ranging from 17 to 50 kg (approximately 37\u2013110 lbs.). A paleoenvironmental study reconstructed the habitat of <em>Proconsul <\/em>to be a dense, closed-canopy tropical forest (Michel Et al. 2014). No caudal vertebrae (tail bones) have been found in direct association with <em>Proconsul <\/em>post-crania, and the morphology of the sacrum is consistent with <em>Proconsul<\/em> lacking a tail (Russo 2016; Ward Et al. 1991).<\/p>\n<p class=\"import-Normal\">Overall, the African ape fossil record in the Late Miocene is sparse, with seven fossil localities dating between eleven and five million years ago (Pickford Et al. 2009). Nevertheless, most species of great apes live in Africa today. Where did the progenitors of modern African apes arise? Did they evolve in Africa or somewhere else? The paucity of apes in the Late Miocene of Africa stands in contrast to the situation in Eurasia. There, ape diversity was high. Furthermore, several Eurasian ape fossils show morphological affinities with modern hominoids (apes). Because of this, some paleoanthropologists suggest that the ancestors of modern African great apes recolonized Africa from Eurasia toward the end of the Miocene (Begun 2002). However, discoveries of Late Miocene hominoids like the Kenyan <em>Nakalipithecus<\/em> (9.9 million to 9.8 million years ago), the Ethiopian <em>Chororapithecus<\/em> (10.7 million to 10.1 million years ago), and the late-Middle Miocene Namibian <em>Otavipithecus<\/em> (13 million to 12 million years ago) fuel an alternative hypothesis\u2014namely that African hominoid diversity was maintained throughout the Miocene and that one of these taxa might, in fact, be the last common ancestor of extant African apes (Kunimatsu Et al. 2007; Mocke Et al. 2002). The previously underappreciated diversity of Late Miocene apes in Africa might be due to poor sampling of the fossil record in Africa.<\/p>\n<h4 class=\"import-Normal\"><em>Eurasian Ape Diversity<\/em><\/h4>\n<p class=\"import-Normal\">With the establishment of the <em>Gomphotherium<\/em> Landbridge (a result of the closure of the Eastern Mediterranean seaway; R\u00f6gl 1999), the Middle Miocene was an exciting time for hominoid radiations outside of Africa (see Figure 9.23). Eurasian hominoid species exploited their environments in many different ways in the Miocene. Food exploitation ranged from soft-fruit feeding in some taxa to hard-object feeding in others, in part owing to seasonal fluctuations and the necessary adoptions of fallback foods (DeMiguel Et al. 2014). For example, the molars of <em>Oreopithecus bambolii<\/em> (Family Hominidae) have relatively long lower-molar shearing crests, suggesting that this hominoid was very folivorous (Ungar and Kay 1995). Associated with variation in diet, there is great variation in the degree to which cranial features (e.g., zygomatic bone or supraorbital tori) are developed across the many taxa (Cameron 1997); however, Middle Miocene fossils tend to exhibit relatively thick molar enamel and relatively robust jaws (Andrews and Martin 1991).<\/p>\n<p class=\"import-Normal\">In Spain, the cranium with upper dentition, part of a mandible, and partial skeleton of <em>Pliobates <\/em>(Family Pliobatidae), a small-bodied ape (4\u20135 kg; 9\u201311 lbs.), was discovered in deposits dating to 11.6 million years ago (Alba Et al. 2015). It is believed to be a frugivore with a relative brain size that overlaps with modern cercopithecoids. The fossilized postcrania of <em>Pliobates<\/em> suggest that this ape might have had a unique style of locomotion, including the tendency to walk across the branches of trees with its palms facing downward and flexible wrists that permitted rotation of the forearm during climbing. However, the anatomy of the distal humerus differs from those of living apes in ways that suggest that <em>Pliobates<\/em> was less efficient at stabilizing its elbow while suspended (Benefit and McCrossin 2015). Two other recently described apes from Spain, <em>Pierolapithecus <\/em>and <em>Anoiapithecus<\/em>, are known from relatively complete skeletons. <em>Pierolapithecus<\/em> had a very projecting face and thick molar enamel as well as some skeletal features that suggest (albeit controversially) a less suspensory locomotor style than in extant apes (Moy\u00e0-Sol\u00e0 Et al. 2004). In contrast to <em>Pierolapithecus<\/em>, the slightly younger <em>Anoiapithecus<\/em> has a very flat face (Moy\u00e0-Sol\u00e0 Et al. 2009).<\/p>\n<p class=\"import-Normal\">Post-cranial evidence for suspensory or well-developed orthograde behaviours in apes does not appear until the Late Miocene of Europe. Primary evidence supporting these specialized locomotor modes includes the relatively short lumbar vertebrae of <em>Oreopithecus <\/em>(Figure 9.26) and <em>Dryopithecus<\/em> (Maclatchy 2004). Further, fossil material of the lower torso of <em>O. bambolii <\/em>(which dates to the <em>Pan<\/em>-hominin divergence) conveys a higher degree of flexion-extension abilities in the lumbar region (lower back) than what is possible in extant apes. Additionally, the hindlimb of <em>O. bambolii <\/em>is suggested to have supported powerful hip adduction during climbing (Hammond Et al. 2020). The Late Miocene saw the extinction of most of the Eurasian hominoids in an event referred to as the Vallesian Crisis (Agust\u00ed Et al. 2003). Among the latest surviving hominoid taxa in Eurasia were <em>Oreopithecus<\/em> and <em>Gigantopithecus<\/em>, the latter of which held out until the Pleistocene in Asia and was probably even sympatric with <em>Homo erectus<\/em> (Cachel 2015).<\/p>\n<figure style=\"width: 436px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image8-2-1.jpg\" alt=\"Posterior view of ancient ape skeleton.\" width=\"436\" height=\"775\" \/><figcaption class=\"wp-caption-text\">Figure 9.26: Skeleton of Oreopithecus bambolii. Credit: <a class=\"rId107\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Oreopithecus_bambolii_1.JPG\">Oreopithecus bambolii 1<\/a> by <a class=\"rId108\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Ghedoghedo\">Ghedoghedo<\/a> is under a <a class=\"rId109\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>The Origins of Extant Apes<\/strong><\/h3>\n<p class=\"import-Normal\">The fossil record of the extant apes is somewhat underwhelming: it ranges from being practically nonexistent for some taxa (e.g., chimpanzees) to being a little better for others (e.g., humans). There are many possible reasons for these differences in fossil abundance, and many are associated with the environmental conditions necessary for the fossilization of bones. One way to understand the evolution of extant apes that is not so dependent on the fossil record is via molecular evolutionary analyses. This can include counting up the differences in the genetic sequence between two closely related species to estimate the amount of time since these species shared a common ancestor. This is called a molecular clock, and it is often calibrated using fossils of known absolute age that stand in for the last common ancestor of a particular clade. Molecular clock estimates have placed the Hylobatidae and Hominidae split between 19.7 million and 24.1 million years ago, the African ape and Asian ape split between 15.7 million and 19.3 million years ago, and the split of Hylobatidae into its current genera between 6.4 million and 8 million years ago (Israfil Et al. 2011).<\/p>\n<h4 class=\"import-Normal\"><em>Small Ape Origins and Fossils<\/em><\/h4>\n<p class=\"import-Normal\">Unfortunately, the fossil record for the small (formerly \u201clesser\u201d) apes is meager, particularly in Miocene deposits. One possible early hylobatid is <em>Laccopithecus robustus<\/em>, a Late Miocene catarrhine from China (Harrison 2016). Although it does share some characteristics with modern gibbons and siamangs (including an overall small body size and a short face), <em>Laccopithecus<\/em> most likely represents a plesiomorphic stem catarrhine and is therefore distantly related to extant apes (Jablonski &amp; Chaplin 2009). A more likely candidate for the hylobatid stem is another Late Miocene taxon from China, <em>Yuanmoupithecus xiaoyuan<\/em>. Interpretation of its phylogenetic standing, however, is complicated by contradicting dental features\u2014some of them quite plesiomorphic\u2014which some believe best place <em>Yuanmoupithecus<\/em> as a stem hylobatid (Harrison 2016). Recently, a Middle Miocene Indian fossil ape, <em>Kapi ramnagarensis<\/em>, has extended the fossil record of small apes by approximately five million years. Its teeth are suggestive of a shift to a more frugivorous diet and it is likely a stem hylobatid (Gilbert Et al. 2020). The history of Hylobatidae becomes clearer in the Pleistocene, with fossils representing extant genera.<\/p>\n<h4 class=\"import-Normal\"><em>Great Ape Origins and Fossils<\/em><\/h4>\n<p class=\"import-Normal\">The most extensive fossil record of a modern great ape is that of our own genus, <em>Homo<\/em>. However, the evolutionary history of the Asian great ape, the orangutan (<em>Pongo<\/em>), is becoming clearer. Today, orangutans are found only on the islands of Borneo and Sumatra. However, Pleistocene-aged teeth, attributed to <em>Pongo<\/em>, have been found in Cambodia, China, Laos, Peninsular Malaysia, and Vietnam\u2014demonstrating the vastness of the orangutan\u2019s previous range (Ibrahim Et al. 2013; Wang Et al. 2014). <em>Sivapithecus <\/em>from the Miocene of India and Pakistan is represented by many specimens, including parts of the face. <em>Sivapithecus<\/em> is very similar to <em>Pongo<\/em>, especially in the face, and it probably is closely related to ancestral orangutans (Pilbeam 1982). Originally, jaws and teeth belonging to the former genus <em>Ramapithecus<\/em> were thought to be important in the origin of humans (Simons 1961), but now these are recognized as specimens of <em>Sivapithecus<\/em> (Kelley 2002). Postcranial bones of <em>Sivapithecus<\/em>, however, suggest a more generalized locomotor mode\u2014including terrestrial locomotion\u2014than seen in <em>Pongo <\/em>(Pilbeam Et al. 1990). Stable carbon and oxygen isotope data from dental enamel have reconstructed the paleoecological space of <em>Sivapithecus <\/em>(as well as the contemporaneous Late Miocene pongine <em>Khoratpithecus<\/em>) within the canopies of forested habitats (Habinger Et al. 2022).<\/p>\n<p class=\"import-Normal\">A probable close relative of <em>Sivapithecus <\/em>is the amazing <em>Gigantopithecus<\/em> (see Figure 9.24). Known only from teeth and jaws from China and India (e.g., Figure 9.27), this ape probably weighed as much as 270 kg (595 lbs.) and was likely the largest primate ever (Bocherens Et al. 2017). Because of unique features of its teeth (including molarized premolars and patterns of wear) and its massive size, it has been reconstructed as a bamboo specialist, somewhat like the modern panda. Small silica particles (phytoliths) from grasses have been found stuck to the molars of <em>Gigantopithecus<\/em> (Ciochon Et al. 1990). Recent studies evaluating the carbon isotope composition of the enamel sampled from <em>Gigantopithecus<\/em> teeth suggest that this ape exploited a wide range of vegetation, including fruits, leaves, roots, and bamboo (Bocherens Et al. 2017). Its face is reminiscent of that of modern orangutans and it might belong in the same family, Pongidae (Kelley 2002).<\/p>\n<figure style=\"width: 488px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image12.jpg\" alt=\"Superior view of mandible and teeth.\" width=\"488\" height=\"533\" \/><figcaption class=\"wp-caption-text\">Figure 9.27: Cast of the mandible of Gigantopithecus blacki. Credit: <a class=\"rId111\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Gigantopithecus%20blacki%20mandible%20010112.jpg\">Gigantopithecus blacki mandible 010112<\/a> by <a class=\"rId112\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Wilson44691\">Wilson44691<\/a> is under a <a class=\"rId113\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">In Africa, the first fossil to be confidently attributed to <em>Pan<\/em>, and known to be the earliest evidence of a chimpanzee, was described based on teeth found in Middle Pleistocene deposits in the Eastern Rift Valley of Kenya (McBrearty and Jablonski 2005). Paleoenvironmental reconstructions of this locality suggest that this early chimpanzee was living in close proximity to early <em>Homo<\/em> in a closed-canopy wooded habitat. Similarly, fossil teeth and mandibular remains attributed to two species of Middle-Late Miocene apes\u2014<em>Chororapithecus abyssinicus<\/em> (from Ethiopia; Suwa Et al. 2007) and <em>Nakalipithecus nakayamai<\/em> (from Kenya; Kunimatsu Et al. 2007)\u2014have been suggested as basal members of the gorilla clade.<\/p>\n<p class=\"import-Normal\">While the deposits of Eastern Africa have yielded a profound record of our fossil hominin ancestors, the continent\u2019s rainforests remain a \u201cpalaeontological desert\u201d (Rosas Et al. 2022). Clearly, more work is needed to fill in the large gaps in the fossil record of the nonhuman great apes. The twentieth century witnessed the discovery of many hominin fossils in East Africa, which have been critical for improving our understanding of human evolution. While twenty-first-century conservationists fight to prevent the extinction of the living great apes, perhaps efforts by twenty-first-century paleoanthropologists will yield the evolutionary story of these, our closest relatives.<\/p>\n<div class=\"textbox shaded\">\n<h2>Summary<\/h2>\n<p>While there are large gaps in the fossil records linking primates to early hominins, evolutionary trends make it clear that humans are one branch of the broader primate family tree. In this chapter we go over the major development that characterize primate evolution: enhanced vision, grasping hands and feet, greater reliance on social behaviour, and increased brain complexity. It is these traits which distinguish primates from other mammals and furthermore, help define major subsections within the primate class, such as strepsirrhines, haplorhines, monkeys, apes, and ultimately hominins.<\/p>\n<p>Within this chapter, we also examine how anthropologists reconstruct these evolutionary relationships. Fossil evidence has provided key information about when and where different primates lived, while genetic data such as skeletal features allow researchers to understand how extinct species moved, ate, and interacted with their environments. Just as crucial are the influences of Earth\u2019s changing environments: continental drift, glacial cycles, and long-term climate shifts have repeatedly reshaped habitats, driving both extinctions and the emergence of new adaptive forms, including the emergence of our own human lineage.<\/p>\n<h2 class=\"import-Normal\">Review Questions<strong><br \/>\n<\/strong><\/h2>\n<ul>\n<li>Compare three major hypotheses about primate origins, making reference to each one\u2019s key ecological reason for primate uniqueness.<\/li>\n<li>Explain how changes in temperature, rainfall, and vegetation led to major changes in primate biogeography over the Early Tertiary.<\/li>\n<li>List some euprimate features that plesiadapiforms have and some that they lack.<\/li>\n<li>Contrast adapoids and omomyoids in terms of life habits.<\/li>\n<li>Describe one piece of evidence for each of the adapoid, omomyoid, and tarsier origin hypotheses for anthropoids.<\/li>\n<li>Discuss the biogeography of the origins of African great apes and orangutans using examples from the Miocene ape fossil record.<\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\">Key Terms<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\"><strong>Adapoidea<\/strong>: Order: Primates. One of the earliest groups of euprimates (true primates; earliest records from the early Eocene).<\/p>\n<p class=\"import-Normal\"><strong>A<\/strong><strong>daptive radiations<\/strong>: Rapid diversifications of single lineages into many species which may present unique morphological features in response to different ecological settings.<\/p>\n<p class=\"import-Normal\"><strong>Ancestral traits<\/strong>: Features that were inherited from a common ancestor and which remain (largely) unchanged.<\/p>\n<p class=\"import-Normal\"><strong>Anthropoids<\/strong>:Group containing monkeys and apes, including humans.<\/p>\n<p class=\"import-Normal\"><strong>Auditory bulla<\/strong>: The rounded bony floor of the middle ear cavity.<\/p>\n<p class=\"import-Normal\"><strong>Bilophodonty<\/strong>: Dental condition in which the cusps of molar teeth form ridges (or lophs) separated from each other by valleys (seen, e.g., in modern catarrhine monkeys).<\/p>\n<p class=\"import-Normal\"><strong>Catarrhines<\/strong>: Order: Primates; Suborder: Anthropoidea; Infraorder: Catarrhini. Group, with origins in Africa and Asia, that contains monkeys and apes, including humans.<\/p>\n<p class=\"import-Normal\"><strong>Clade<\/strong>:Group containing all of the descendants of a single ancestor. A portion of a phylogenetic tree represented as a bifurcation (node) in a lineage and all of the branches leading forward in time from that bifurcation.<\/p>\n<p class=\"import-Normal\"><strong>Convergent evolution<\/strong>: The independent evolution of a morphological feature in animals not closely related (e.g., wings in birds and bats).<\/p>\n<p class=\"import-Normal\"><strong>Crown<\/strong>: Smallest monophyletic group (clade) containing a specified set of extant taxa and all descendants of their last common ancestor.<\/p>\n<p class=\"import-Normal\"><strong>Diastema<\/strong>: Space between adjacent teeth.<\/p>\n<p class=\"import-Normal\"><strong>Diffuse coevolution<\/strong>: The ecological interaction between whole groups of species (e.g., primates) with whole groups of other species (e.g., fruiting trees).<\/p>\n<p class=\"import-Normal\"><strong>Ectotympanic<\/strong>: Bony ring or tube that holds the tympanic membrane (eardrum).<\/p>\n<p class=\"import-Normal\"><strong>Euprimates<\/strong>: Order: Primates. True primates or primates of modern aspect.<\/p>\n<p class=\"import-Normal\"><strong>Haplorhines<\/strong>: Group containing catarrhines, platyrrhines, and tarsiers.<\/p>\n<p class=\"import-Normal\"><strong>Hominins<\/strong>: Modern humans and any extinct relatives more closely related to us than to chimpanzees.<\/p>\n<p class=\"import-Normal\"><strong>Mandibular symphysis<\/strong>: Fibrocartilaginous joint between the left and right mandibular segments, located in the midline of the body.<\/p>\n<p class=\"import-Normal\"><strong>Omomyoidea<\/strong>: Order: Primates; Superfamily: Omomyoidea. One of the earliest groups of euprimates (true primates; earliest record in the early Eocene).<\/p>\n<p class=\"import-Normal\"><strong>Petrosal bone<\/strong>: The portion of the temporal bone that houses the inner ear apparatus.<\/p>\n<p class=\"import-Normal\"><strong>Plagiaulacoid<\/strong>: Dental condition where at least one of the lower cheek-teeth (molars or premolars) is a laterally compressed blade.<\/p>\n<p class=\"import-Normal\"><strong>Platyrrhines<\/strong>: Order: Primates; Suborder: Anthropoidea; Infraorder: Platyrrhini. Group containing monkeys found in the Americas.<\/p>\n<p class=\"import-Normal\"><strong>Plesiadapiforms<\/strong>: Order: Plesiadapiformes. Archaic primates or primate-like placental mammals (Early Paleocene\u2013Late Eocene).<\/p>\n<p class=\"import-Normal\"><strong>P<\/strong><strong>lesiomorphic<\/strong>: Having features that are shared by different groups which arose from a common ancestor.<\/p>\n<p class=\"import-Normal\"><strong>Stem<\/strong>: Taxa that are basal to a given crown group but are more closely related to the crown group than to the closest living sister taxon of the crown group.<\/p>\n<p class=\"import-Normal\"><strong>Strepsirrhines<\/strong>: Order: Primates; Suborder: Stresirrhini. Group containing lemurs, lorises, and galagos (does not include tarsiers).<\/p>\n<p class=\"import-Normal\"><strong>Toothcomb<\/strong>: Dental condition found in modern strepsirrhines in which the lower incisors and canines are laterally compressed and protrude forward at a nearly horizontal inclination. This structure is used in grooming.<\/p>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<h2 class=\"import-Normal\">For Further Exploration<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\">Beard, Chris. 2004. <em>The Hunt for the Dawn Monkey: Unearthing the Origins of Monkeys, Apes, and Humans<\/em>. Berkeley: University of California Press.<\/p>\n<p class=\"import-Normal\">Begun, David R. 2010. \u201cMiocene Hominids and the Origins of the African Apes and Humans.\u201d <em>Annual Review of Anthropology<\/em> 39: 67\u201384.<\/p>\n<p class=\"import-Normal\">Fleagle, John G. 2013. <em>Primate Adaptation and Evolution.<\/em> Third edition. San Diego, CA: Academic Press.<\/p>\n<p class=\"import-Normal\">Gebo, Daniel L., ed. 1993. <em>Postcranial Adaptations in Nonhuman Primates<\/em>. Dekalb: Northern Illinois University Press.<\/p>\n<p class=\"import-Normal\">Godfrey, Laurie R., and William L. Jungers. 2002. \u201cQuaternary Fossil Lemurs.\u201d In <em>The Primate Fossil Record, <\/em>edited by Walter C. Hartwig, 97\u2013121. Cambridge: Cambridge University Press.<\/p>\n<p class=\"import-Normal\">Godinot, Marc. 2006. \u201cLemuriform Origins as Viewed from the Fossil Record.\u201d <em>Folia Primatologica<\/em> 77 (6): 446\u2013464.<\/p>\n<p class=\"import-Normal\">Kay, Richard F. 2018. \u201c100 Years of Primate Paleontology.\u201d <em>American Journal of Physical Anthropology<\/em> 165 (4): 652\u2013676.<\/p>\n<p class=\"import-Normal\">Marivaux, Laurent. 2006. \u201cThe Eosimiid and Amphipithecid Primates (Anthropoidea) from the Oligocene of the Bugti Hills (Balochistan, Pakistan): New Insight into Early Higher Primate Evolution in South Asia.\u201d <em>Palaeovertebrata, Montpellier <\/em>34 (1\u20132): 29\u2013109.<\/p>\n<p class=\"import-Normal\">Martin, R. D. 1990. <em>Primate Origins and Evolution<\/em><em>: A <\/em><em>Phylogenetic Reconstruction<\/em>. Princeton: Princeton University Press.<\/p>\n<p class=\"import-Normal\">Rose, Kenneth D., Marc Godinot, and Thomas M. Bown. 1994. \u201cThe Early Radiation of Euprimates and the Initial Diversification of Omomyidae.\u201d In <em>Anthropoid Origins: The Fossil Evidence, <\/em>edited by John G. Fleagle and Richard F. Kay, 1\u201328. New York: Plenum Press.<\/p>\n<p class=\"import-Normal\">Ross, Callum F. 1999. \u201cHow to Carry Out Functional Morphology.\u201d <em>Evolutionary Anthropology<\/em> 7 (6): 217\u2013222.<\/p>\n<p class=\"import-Normal\">Seiffert, Erik R. 2012. \u201cEarly Primate Evolution in Afro-Arabia.\u201d Evolutionary Anthropology: Issues, News, and Reviews 21(6): 239\u2013253.<\/p>\n<p class=\"import-Normal\">Szalay, Frederic S., and Eric Delson. 1979. Evolutionary History of the Primates. New York: Academic Press.<\/p>\n<p class=\"import-Normal\">Ungar, Peter S. 2002. \u201cReconstructing the Diets of Fossil Primates.\u201d In <em>Reconstructing Behavior in the Primate Fossil Record<\/em>, edited by Joseph Plavcan, Richard F. Kay, William Jungers, and Carel P. van Schaik, 261\u2013296. New York: Kluwer Academic\/Plenum Publishers.<\/p>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\">Agust\u00ed, J., A. Sanz de Siria, and M. Garc\u00e9s M. 2003. \u201cExplaining the End of the Hominoid Experiment in Europe.\u201d <em>Journal of Human Evolution<\/em> 45 (2): 145\u2013153.<\/p>\n<p class=\"import-Normal\">Alba, David M., Sergio Alm\u00e9cija, Daniel DeMiguel, Josep Fortuny, Miriam P\u00e9rez de los R\u00edos, Marta Pina, Josep M. 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Beth Shook, Lara Braff, Katie Nelson, and Kelsie Aguilera, for their time and commitment to making this knowledge freely accessible to all, and for giving us the opportunity to participate in this important project.<\/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_1683_1686\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1686\"><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<h6>Student contributors to this chapter: Angela Durastanti, Bryce Muller, Gabriel Barr, Maisie Babbington-Bolduc<\/h6>\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\">Learning Objectives<\/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_1683_1291\">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_1683_1326\">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_1683_1327\">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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1328\">fossils<\/a><\/strong>, 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\">Within this conversation, naturalists and early <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1329\">paleoanthropologists<\/a><\/strong> (people who study human evolution) speculated about which human traits came first. These included the evolution of a big brain (<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1330\">encephalization<\/a><\/strong>), 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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1331\">morphology<\/a><\/strong> (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_1683_1332\"><strong>East African Rift System<\/strong>(EARS)<\/a>; that is, in Ethiopia, Kenya, Tanzania, and Malawi; see Figure 10.1), 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/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 10.1: 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_1683_1333\">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_1683_1335\"><strong>taxonomy<\/strong><\/a> was primarily based on morphology. Today it is tied to known relationships based on molecular <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1336\">phylogeny<\/a><\/strong> (e.g., based on DNA) or a combination of the two. This is complicated when applied to living <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1334\">taxa<\/a><\/strong>, 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_1683_1337\">cladistics<\/a> <\/strong>and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1338\">phylogenetics<\/a><\/strong><strong>. <\/strong>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_1683_1254\">derived traits<\/a><\/strong>. 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_1683_1249\">clade<\/a> <\/strong>(Figure 10.2). For example, if we use feathers as a trait, we can group pigeons and ostriches into the clade of birds. 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 10.2: 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<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1339\">lumpers<\/a><\/strong>,\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<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1340\">splitters<\/a><\/strong>,\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_1683_1341\">polytypic<\/a><\/strong>. 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 10.3: 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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1342\">chronospecies<\/a><\/strong> (Figure 10.3). 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. Large-scale changes in global and regional climate, as well as alterations to the environment, are thought to be linked to all hominin diversification, dispersal, and extinction (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_1683_1345\">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_1683_1343\">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_1683_1344\">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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1138\">isotopes<\/a><\/strong>; 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 10.4). 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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1347\">flora<\/a><\/strong> 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 10.4: 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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1348\">arboreal<\/a><\/strong> lifestyle (one living in trees) to a terrestrial one where bipedalism was a more efficient form of locomotion (Figure 10.5). 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_1683_1350\">Aridity Hypothesis<\/a><\/strong>. This hypothesis states that the long-term <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1349\">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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image46.png\" alt=\"Photograph showing a dry, open savannah environment.\" width=\"647\" height=\"486\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 10.5: 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, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1354\">ungulates<\/a><\/strong> (hoofed mammals of various kinds) that had generalized diets fared better than those with specialized diets (Vrba 1988, 1998). <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1352\">Specialist<\/a><\/strong> eaters faced extinction at greater rates than their <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1351\">generalist<\/a> <\/strong>counterparts because they were unable to adapt to new environments (Vrba 2000). Thus, periods with extreme climate change would be associated with high <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1353\">faunal turnover<\/a><\/strong>: 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_1683_1391\">Quaternary Ice Age<\/a><\/strong>, between 2.5 mya and 3 mya, brought extreme global, cyclical <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1355\">interglacial<\/a><\/strong> and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1356\">glacial<\/a><\/strong> 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 behavioural 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 behavioural 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<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_1683_1357\">obligate bipedalism<\/a><\/strong>, is important in distinguishing our species from the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1358\">extant<\/a><\/strong> (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 theories:<\/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 centre 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 10.6: 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 10.6). 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 behaviours. These adaptations occur throughout the skeleton and are summarized in Figure 10.7.<\/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_1683_1336\"><strong>postcranium<\/strong><\/a> and are outlined in Figure 10.7. 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 centre 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_1683_1392\">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_1683_1360\">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><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<p><span style=\"color: #000000\">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<table class=\"no-lines landscape alignleft\" style=\"border-collapse: collapse;width: 98.92%;height: 54px\" border=\"0\">\n<tbody>\n<tr style=\"height: 67px\">\n<td style=\"width: 100%;height: 54px\">Figure 10.7: Skeletal comparisons between modern humans (obligate bipeds) and nonobligate bipeds (e.g., chimpanzees). Credit:<a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\"> Skeletal comparisons between modern humans and nonobligate bipeds (Figure 9.6)<\/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>.<\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<div align=\"left\">\n<table class=\"grid\">\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 centre 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<div class=\"textbox shaded\" style=\"background: var(--lightblue)\">\n<h2>Special Topic: Fear of Snakes \u2014 A Cultural or Biological Adaptation?<\/h2>\n<figure id=\"attachment_680\" aria-describedby=\"caption-attachment-680\" style=\"width: 393px\" class=\"wp-caption alignright\"><img class=\"wp-image-680\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2023\/06\/snake-2319873_1920.jpg\" alt=\"\" width=\"393\" height=\"262\" \/><figcaption id=\"caption-attachment-680\" class=\"wp-caption-text\">Figure 10.8: A constricting snake (adder) with patterned scales and opened mouth. Credit: <em data-start=\"542\" data-end=\"582\">Snake, Adder, Serpent, Reptile, Animal<\/em> is free to use under the Pixabay <a href=\"https:\/\/pixabay.com\/service\/license-summary\/\">Content License.<\/a><\/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 responsible for visual attention and oculomotor behaviour (Isbell, L., 2009).<\/p>\n<figure style=\"width: 316px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/upload.wikimedia.org\/wikipedia\/commons\/thumb\/9\/96\/Ra_slays_Apep_%28tomb_scene_in_Deir_el-Medina%29%28improved_contrast%29.png\/250px-Ra_slays_Apep_%28tomb_scene_in_Deir_el-Medina%29%28improved_contrast%29.png\" alt=\"File:Ra slays Apep (tomb scene in Deir el-Medina)(improved contrast).png\" width=\"316\" height=\"236\" \/><figcaption class=\"wp-caption-text\">Figure 10.9: Tomb scene depicting the serpent god Apep (Apophis) wounded by the cat god Miuty, alternatively the sun god Ra in cat form, from a tomb in Deir el-Medina, Luxor, Egypt. Credit: <em data-start=\"265\" data-end=\"331\">Ra slays Apep (tomb scene in Deir el-Medina) (improved contrast)<\/em> by Hajor is under <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/deed.en\">Creative Commons Attribution-Share Alike 3.0<\/a><\/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 behaviours toward threats like snakes. This blend of instinct and cultural influence is not only seen in behaviour 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 behaviours 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 behaviour.<\/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 10.10; 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. This might imply that, over evolutionary time, the need for display and dominance among males has reduced, as has our sexual dimorphism. 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_304\" aria-describedby=\"caption-attachment-304\" style=\"width: 640px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-288\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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-304\" class=\"wp-caption-text\">Figure 10.10: 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 behaviours (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_1683_1361\">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_1683_1393\">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 10.11). 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_304-2\" aria-describedby=\"caption-attachment-304-2\" style=\"width: 706px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-289\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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-304-2\" class=\"wp-caption-text\">Figure 10.11a 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<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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1362\">frugivorous<\/a><\/strong> (fruit-eating) chimpanzee or the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1363\">folivorous<\/a><\/strong> (foliage-eating) gorilla, as discussed in Chapter 6.<\/span><\/p>\n<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_1683_1364\">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. 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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1365\">dental formula<\/a><\/strong>. 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_1683_1271\">incisors<\/a><\/strong>; the pointy <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1274\">canines<\/a><\/strong>; the small, flatter <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1366\">premolars<\/a><\/strong>; and the larger hind <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1273\">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<figure style=\"width: 380px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 10.12: 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_1683_1389\">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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1367\">procumbent<\/a><\/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\">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_1683_1390\">incisiform<\/a> <\/strong>(Figure 10.12). 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 dominance.<\/span><\/p>\n<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. This might suggest that, over evolutionary time, the need for display and dominance among males has reduced, as has our sexual dimorphism. 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_1683_1277\">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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1368\">honing P3<\/a><\/strong>, 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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1369\">parabolic<\/a><\/strong> (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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1370\">enamel<\/a><\/strong> 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_1683_1279\">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 behaviours 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<div class=\"textbox\">\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_1683_1371\">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 10.13). 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_304-3\" aria-describedby=\"caption-attachment-304-3\" style=\"width: 579px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-291 \" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/9.11.jpg\" alt=\"Four views of an ancient skull are shown on a black background.\" width=\"579\" height=\"579\" \/><figcaption id=\"caption-attachment-304-3\" class=\"wp-caption-text\">Figure 10.13: 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<\/div>\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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 10.14: 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 10.14).<\/span><\/p>\n<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_1683_1394\">gracile<\/a> <\/strong>australopithecines\u201d because of their smaller and less robust features compared to the divergent \u201c<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1395\">robust<\/a><\/strong>\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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1372\">volcanic tufts<\/a><\/strong> that have accumulated over millions of years. These tufts may be dated using absolute dating techniques, such as Potassium-Argon dating (described in Chapter 8). This means that it is possible to know a relatively refined date for any fossil if the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1373\">context<\/a> <\/strong>of that find is known. 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 10.15: 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 10.15). 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 10.16). 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 8; Hay and Leakey 1982; Leakey and Hay 1979) have drawn much attention.<\/span><\/p>\n<figure id=\"attachment_304-4\" aria-describedby=\"caption-attachment-304-4\" style=\"width: 643px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-294 \" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/9.14.jpg\" alt=\"Two images of life-like reconstructions of female and male Au. afarensis.\" width=\"643\" height=\"322\" \/><figcaption id=\"caption-attachment-304-4\" class=\"wp-caption-text\">Figure 10.16 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image40.png\" alt=\"A partial skeleton includes bones of the cranium, mandible, and postcranium.\" width=\"320\" height=\"772\" \/><figcaption class=\"wp-caption-text\">Figure 10.17: 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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1374\">prognathic<\/a><\/strong> 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 10.17). Further evidence of bipedalism is seen in the Laetoli Footprints, which are associated with <em data-start=\"92\" data-end=\"107\">Au. afarensis<\/em> (see Chapter 8). <\/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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1129\">relative dating<\/a><\/strong>, 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 10.18). 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<figure style=\"width: 570px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 10.18: 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 10.19). 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<figure id=\"attachment_304-5\" aria-describedby=\"caption-attachment-304-5\" style=\"width: 548px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-297 \" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/9.17.jpg\" alt=\"Four views of an ancient skull are shown on a black background.\" width=\"548\" height=\"548\" \/><figcaption id=\"caption-attachment-304-5\" class=\"wp-caption-text\">Figure 10.19: 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 10.20; 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 10.20: 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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1376\">fallback foods<\/a><\/strong> 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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1377\">monophyletic<\/a><\/strong>) 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 10.21; 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<figure id=\"attachment_304-6\" aria-describedby=\"caption-attachment-304-6\" style=\"width: 666px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-299 \" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/9.19.jpg\" alt=\"Five views of a beige partial skull on a black background.\" width=\"666\" height=\"444\" \/><figcaption id=\"caption-attachment-304-6\" class=\"wp-caption-text\">Figure 10.21: 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 10.22 and 10.23; 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_1683_1378\">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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1379\">hyper-robust<\/a><\/strong>\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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image14-1.png\" alt=\"Life-like reconstruction of Paranthropus boisei.\" width=\"557\" height=\"557\" \/><figcaption class=\"wp-caption-text\">Figure 10.22: 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_304-7\" aria-describedby=\"caption-attachment-304-7\" style=\"width: 565px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-301 \" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/9.21.jpg\" alt=\"Three views of an ancient skull are shown on a black background.\" width=\"565\" height=\"565\" \/><figcaption id=\"caption-attachment-304-7\" class=\"wp-caption-text\">Figure 10.23: \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 10.24; 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_304-8\" aria-describedby=\"caption-attachment-304-8\" style=\"width: 572px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-302 \" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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-304-8\" class=\"wp-caption-text\">Figure 10.24: 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_303\" aria-describedby=\"caption-attachment-303\" style=\"width: 570px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-303 \" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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-303\" class=\"wp-caption-text\">Figure 10.25: 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><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 10.25; 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<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_1683_1380\">Early Stone Age (ESA)<\/a><\/strong> marks the beginning of recognizable technology made by our human ancestors. Stone-tool (or <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1381\">lithic<\/a><\/strong>) technology is defined by the fracturing of rocks and the manufacture of tools through a process called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1382\">knapping<\/a><\/strong>. 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_1683_1383\">techno-complex<\/a><\/strong>, a term encompassing multiple <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1384\">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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1386\">knappers<\/a><\/strong>) began to change the ways they detached <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1385\">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. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1387\">Large Cutting Tools (LCTs) <\/a><\/strong> 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image29-1.png\" alt=\"A technical line drawing of an Oldowan chopper.\" width=\"324\" height=\"275\" \/><figcaption class=\"wp-caption-text\">Figure 10.26: 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_1683_1388\">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 10.26).<\/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\">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 metres tall and females at 1.24 metres.<\/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>Summary<\/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\"><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 centre of gravity (i.e., between the feet) in bipeds.<\/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. 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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> 12 (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\">Van Le, Q., Isbell, L. A., Matsumoto, J., Nguyen, M., Hori, E., Maior, R. S., Tomaz, C., Tran, A. H., Ono, T., &amp; Nishijo, H. (2013). Pulvinar neurons reveal neurobiological evidence of past selection for rapid detection of snakes. <em>Proceedings of the National Academy of Sciences, 110<\/em>(47), 19000\u201319005. <\/span><\/p>\n<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_1683_1685\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1685\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<div class=\"textbox\">\n<h6>Student contributors to this chapter: Angela Durastanti, Bryce Muller, Gabriel Barr, Maisie Babbington-Bolduc<\/h6>\n<p>Learn more about <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/about-the-authors\/\">authors and editors<\/a>,<a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/student-contributors\/\"> Hess' student contributions<\/a>, and <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/versioning-history\/\">versioning history<\/a>.<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\">Learning Objectives<\/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_1683_1291\">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_1683_1326\">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_1683_1327\">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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1328\">fossils<\/a><\/strong>, 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\">Within this conversation, naturalists and early <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1329\">paleoanthropologists<\/a><\/strong> (people who study human evolution) speculated about which human traits came first. These included the evolution of a big brain (<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1330\">encephalization<\/a><\/strong>), 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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1331\">morphology<\/a><\/strong> (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_1683_1332\"><strong>East African Rift System<\/strong>(EARS)<\/a>; that is, in Ethiopia, Kenya, Tanzania, and Malawi; see Figure 10.1), 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/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 10.1: 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_1683_1333\">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_1683_1335\"><strong>taxonomy<\/strong><\/a> was primarily based on morphology. Today it is tied to known relationships based on molecular <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1336\">phylogeny<\/a><\/strong> (e.g., based on DNA) or a combination of the two. This is complicated when applied to living <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1334\">taxa<\/a><\/strong>, 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_1683_1337\">cladistics<\/a> <\/strong>and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1338\">phylogenetics<\/a><\/strong><strong>. <\/strong>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_1683_1254\">derived traits<\/a><\/strong>. 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_1683_1249\">clade<\/a> <\/strong>(Figure 10.2). For example, if we use feathers as a trait, we can group pigeons and ostriches into the clade of birds. 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 10.2: 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<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1339\">lumpers<\/a><\/strong>,\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<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1340\">splitters<\/a><\/strong>,\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_1683_1341\">polytypic<\/a><\/strong>. 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 10.3: 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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1342\">chronospecies<\/a><\/strong> (Figure 10.3). 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. Large-scale changes in global and regional climate, as well as alterations to the environment, are thought to be linked to all hominin diversification, dispersal, and extinction (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_1683_1345\">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_1683_1343\">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_1683_1344\">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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1138\">isotopes<\/a><\/strong>; 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 10.4). 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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1347\">flora<\/a><\/strong> 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 10.4: 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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1348\">arboreal<\/a><\/strong> lifestyle (one living in trees) to a terrestrial one where bipedalism was a more efficient form of locomotion (Figure 10.5). 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_1683_1350\">Aridity Hypothesis<\/a><\/strong>. This hypothesis states that the long-term <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1349\">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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image46.png\" alt=\"Photograph showing a dry, open savannah environment.\" width=\"647\" height=\"486\" \/><figcaption class=\"wp-caption-text\"><span style=\"color: #000000\">Figure 10.5: 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, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1354\">ungulates<\/a><\/strong> (hoofed mammals of various kinds) that had generalized diets fared better than those with specialized diets (Vrba 1988, 1998). <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1352\">Specialist<\/a><\/strong> eaters faced extinction at greater rates than their <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1351\">generalist<\/a> <\/strong>counterparts because they were unable to adapt to new environments (Vrba 2000). Thus, periods with extreme climate change would be associated with high <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1353\">faunal turnover<\/a><\/strong>: 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_1683_1391\">Quaternary Ice Age<\/a><\/strong>, between 2.5 mya and 3 mya, brought extreme global, cyclical <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1355\">interglacial<\/a><\/strong> and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1356\">glacial<\/a><\/strong> 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 behavioural 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 behavioural 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<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_1683_1357\">obligate bipedalism<\/a><\/strong>, is important in distinguishing our species from the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1358\">extant<\/a><\/strong> (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 theories:<\/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 centre 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 10.6: 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 10.6). 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 behaviours. These adaptations occur throughout the skeleton and are summarized in Figure 10.7.<\/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_1683_1336\"><strong>postcranium<\/strong><\/a> and are outlined in Figure 10.7. 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 centre 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_1683_1392\">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_1683_1360\">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><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<p><span style=\"color: #000000\">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<table class=\"no-lines landscape alignleft\" style=\"border-collapse: collapse;width: 98.92%;height: 54px\" border=\"0\">\n<tbody>\n<tr style=\"height: 67px\">\n<td style=\"width: 100%;height: 54px\">Figure 10.7: Skeletal comparisons between modern humans (obligate bipeds) and nonobligate bipeds (e.g., chimpanzees). Credit:<a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/chapter-9-early-hominins-2\/\"> Skeletal comparisons between modern humans and nonobligate bipeds (Figure 9.6)<\/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>.<\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<div align=\"left\">\n<table class=\"grid\">\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 centre 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<div class=\"textbox shaded\" style=\"background: var(--lightblue)\">\n<h2>Special Topic: Fear of Snakes \u2014 A Cultural or Biological Adaptation?<\/h2>\n<figure id=\"attachment_680-2\" aria-describedby=\"caption-attachment-680-2\" style=\"width: 393px\" class=\"wp-caption alignright\"><img class=\"wp-image-680\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2023\/06\/snake-2319873_1920.jpg\" alt=\"\" width=\"393\" height=\"262\" \/><figcaption id=\"caption-attachment-680-2\" class=\"wp-caption-text\">Figure 10.8: A constricting snake (adder) with patterned scales and opened mouth. Credit: <em data-start=\"542\" data-end=\"582\">Snake, Adder, Serpent, Reptile, Animal<\/em> is free to use under the Pixabay <a href=\"https:\/\/pixabay.com\/service\/license-summary\/\">Content License.<\/a><\/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 responsible for visual attention and oculomotor behaviour (Isbell, L., 2009).<\/p>\n<figure style=\"width: 316px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/upload.wikimedia.org\/wikipedia\/commons\/thumb\/9\/96\/Ra_slays_Apep_%28tomb_scene_in_Deir_el-Medina%29%28improved_contrast%29.png\/250px-Ra_slays_Apep_%28tomb_scene_in_Deir_el-Medina%29%28improved_contrast%29.png\" alt=\"File:Ra slays Apep (tomb scene in Deir el-Medina)(improved contrast).png\" width=\"316\" height=\"236\" \/><figcaption class=\"wp-caption-text\">Figure 10.9: Tomb scene depicting the serpent god Apep (Apophis) wounded by the cat god Miuty, alternatively the sun god Ra in cat form, from a tomb in Deir el-Medina, Luxor, Egypt. Credit: <em data-start=\"265\" data-end=\"331\">Ra slays Apep (tomb scene in Deir el-Medina) (improved contrast)<\/em> by Hajor is under <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/deed.en\">Creative Commons Attribution-Share Alike 3.0<\/a><\/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 behaviours toward threats like snakes. This blend of instinct and cultural influence is not only seen in behaviour 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 behaviours 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 behaviour.<\/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 10.10; 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. This might imply that, over evolutionary time, the need for display and dominance among males has reduced, as has our sexual dimorphism. 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_304-9\" aria-describedby=\"caption-attachment-304-9\" style=\"width: 640px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-288\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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-304-9\" class=\"wp-caption-text\">Figure 10.10: 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 behaviours (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_1683_1361\">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_1683_1393\">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 10.11). 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_304-10\" aria-describedby=\"caption-attachment-304-10\" style=\"width: 706px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-289\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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-304-10\" class=\"wp-caption-text\">Figure 10.11a 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<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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1362\">frugivorous<\/a><\/strong> (fruit-eating) chimpanzee or the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1363\">folivorous<\/a><\/strong> (foliage-eating) gorilla, as discussed in Chapter 6.<\/span><\/p>\n<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_1683_1364\">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. 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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1365\">dental formula<\/a><\/strong>. 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_1683_1271\">incisors<\/a><\/strong>; the pointy <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1274\">canines<\/a><\/strong>; the small, flatter <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1366\">premolars<\/a><\/strong>; and the larger hind <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1273\">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<figure style=\"width: 380px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 10.12: 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_1683_1389\">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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1367\">procumbent<\/a><\/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\">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_1683_1390\">incisiform<\/a> <\/strong>(Figure 10.12). 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 dominance.<\/span><\/p>\n<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. This might suggest that, over evolutionary time, the need for display and dominance among males has reduced, as has our sexual dimorphism. 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_1683_1277\">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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1368\">honing P3<\/a><\/strong>, 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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1369\">parabolic<\/a><\/strong> (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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1370\">enamel<\/a><\/strong> 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_1683_1279\">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 behaviours 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<div class=\"textbox\">\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_1683_1371\">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 10.13). 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_304-11\" aria-describedby=\"caption-attachment-304-11\" style=\"width: 579px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-291 \" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/9.11.jpg\" alt=\"Four views of an ancient skull are shown on a black background.\" width=\"579\" height=\"579\" \/><figcaption id=\"caption-attachment-304-11\" class=\"wp-caption-text\">Figure 10.13: 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<\/div>\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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 10.14: 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 10.14).<\/span><\/p>\n<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_1683_1394\">gracile<\/a> <\/strong>australopithecines\u201d because of their smaller and less robust features compared to the divergent \u201c<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1395\">robust<\/a><\/strong>\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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1372\">volcanic tufts<\/a><\/strong> that have accumulated over millions of years. These tufts may be dated using absolute dating techniques, such as Potassium-Argon dating (described in Chapter 8). This means that it is possible to know a relatively refined date for any fossil if the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1373\">context<\/a> <\/strong>of that find is known. 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 10.15: 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 10.15). 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 10.16). 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 8; Hay and Leakey 1982; Leakey and Hay 1979) have drawn much attention.<\/span><\/p>\n<figure id=\"attachment_304-12\" aria-describedby=\"caption-attachment-304-12\" style=\"width: 643px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-294 \" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/9.14.jpg\" alt=\"Two images of life-like reconstructions of female and male Au. afarensis.\" width=\"643\" height=\"322\" \/><figcaption id=\"caption-attachment-304-12\" class=\"wp-caption-text\">Figure 10.16 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image40.png\" alt=\"A partial skeleton includes bones of the cranium, mandible, and postcranium.\" width=\"320\" height=\"772\" \/><figcaption class=\"wp-caption-text\">Figure 10.17: 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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1374\">prognathic<\/a><\/strong> 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 10.17). Further evidence of bipedalism is seen in the Laetoli Footprints, which are associated with <em data-start=\"92\" data-end=\"107\">Au. afarensis<\/em> (see Chapter 8). <\/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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1129\">relative dating<\/a><\/strong>, 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 10.18). 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<figure style=\"width: 570px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 10.18: 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 10.19). 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<figure id=\"attachment_304-13\" aria-describedby=\"caption-attachment-304-13\" style=\"width: 548px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-297 \" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/9.17.jpg\" alt=\"Four views of an ancient skull are shown on a black background.\" width=\"548\" height=\"548\" \/><figcaption id=\"caption-attachment-304-13\" class=\"wp-caption-text\">Figure 10.19: 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 10.20; 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 10.20: 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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1376\">fallback foods<\/a><\/strong> 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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1377\">monophyletic<\/a><\/strong>) 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 10.21; 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<figure id=\"attachment_304-14\" aria-describedby=\"caption-attachment-304-14\" style=\"width: 666px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-299 \" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/9.19.jpg\" alt=\"Five views of a beige partial skull on a black background.\" width=\"666\" height=\"444\" \/><figcaption id=\"caption-attachment-304-14\" class=\"wp-caption-text\">Figure 10.21: 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 10.22 and 10.23; 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_1683_1378\">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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1379\">hyper-robust<\/a><\/strong>\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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image14-1.png\" alt=\"Life-like reconstruction of Paranthropus boisei.\" width=\"557\" height=\"557\" \/><figcaption class=\"wp-caption-text\">Figure 10.22: 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_304-15\" aria-describedby=\"caption-attachment-304-15\" style=\"width: 565px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-301 \" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/9.21.jpg\" alt=\"Three views of an ancient skull are shown on a black background.\" width=\"565\" height=\"565\" \/><figcaption id=\"caption-attachment-304-15\" class=\"wp-caption-text\">Figure 10.23: \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 10.24; 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_304-16\" aria-describedby=\"caption-attachment-304-16\" style=\"width: 572px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-302 \" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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-304-16\" class=\"wp-caption-text\">Figure 10.24: 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_303-2\" aria-describedby=\"caption-attachment-303-2\" style=\"width: 570px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-303 \" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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-303-2\" class=\"wp-caption-text\">Figure 10.25: 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><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 10.25; 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<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_1683_1380\">Early Stone Age (ESA)<\/a><\/strong> marks the beginning of recognizable technology made by our human ancestors. Stone-tool (or <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1381\">lithic<\/a><\/strong>) technology is defined by the fracturing of rocks and the manufacture of tools through a process called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1382\">knapping<\/a><\/strong>. 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_1683_1383\">techno-complex<\/a><\/strong>, a term encompassing multiple <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1384\">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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1386\">knappers<\/a><\/strong>) began to change the ways they detached <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1385\">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. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1387\">Large Cutting Tools (LCTs) <\/a><\/strong> 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image29-1.png\" alt=\"A technical line drawing of an Oldowan chopper.\" width=\"324\" height=\"275\" \/><figcaption class=\"wp-caption-text\">Figure 10.26: 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_1683_1388\">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 10.26).<\/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>Cultural Connections and Contemporary Implications<\/h2>\n<p>Evolutionary theories of bipedalism continue to be widely debated. As discussed above, numerous (~30) hypotheses have been proposed to explain why early hominins began walking on two legs. Some remain dominant in the field, while others have been challenged or refined over time. For instance, the <strong>Savannah Hypothesis<\/strong>, which proposed that bipedalism emerged as an adaptation to open grassland environments, has been contested by the <strong>Shore Dweller Hypothesis<\/strong>. Increasing fossil evidence suggests that early hominins may have lived near rivers and wetlands, rather than exclusively in savannahs. This indicates that bipedalism evolved not in hot, dry environments, but rather in more humid, forested regions, where mobility across varied terrain offered selective advantages.<\/p>\n<p>Across these different perspectives, researchers have proposed both biological and cultural explanations for the origins of bipedalism. Lovejoy\u2019s <strong>Male Provisioning Hypothesis,<\/strong> for example, argues that bipedalism developed as males began carrying resources, such as food, to dependent females and offspring. Similarly, other models emphasize the importance of holding and hunting, suggesting that an upright posture freed the arms to transport tools and other objects, which could then be crafted and used more efficiently.<\/p>\n<p>One explanation related to humid forest environments is Niemitz\u2019s <strong>Amphibian Generalist Theory<\/strong> (2010), which proposes that early hominins living near water benefited from plentiful year-round food sources. Plus, moving through water reduces body weight and joint pressure, which may have created selective pressures favouring upright locomotion. This behaviour is still observed in other primates; many adopt bipedal postures in shallow water, and some even swim!<\/p>\n<p>Beyond bipedalism, early hominins experienced a mosaic of evolutionary changes, including alterations to bones, teeth, and, notably, the brain. The brain offers crucial insights into our evolutionary history. Fossil evidence shows a gradual increase in brain size among early hominins (see next chapter on early <em>Homo<\/em>), suggesting significant biological and behavioural shifts. Because the brain is an energy-intensive organ, consuming nearly 18% of the body\u2019s energy, its expansion likely required compensatory adaptations elsewhere. One such theory, the <strong>Expensive Tissue Hypothesis <\/strong>(Aiello &amp; Wheeler, 1995), suggests that as the brain grew, the gut decreased in size to offset the brain's energy demands. This change may have been supported by dietary shifts, not only increased meat consumption (scavenging and opportunistic hunting), but also forms of food processing and the introduction of cooking, making nutrients more accessible and possibly requiring less gut processing time (Wrangham, 1999, 2009).<\/p>\n<p>Brain expansion also had profound implications for childbirth and social behaviour (Rosenberg, 1992). The pelvic modifications required for bipedalism made childbirth more difficult, possibly necessitating assistance from others, and an early form of cooperative care (Burkart et al. 2025). Furthermore, as brain size increased, the capacity for complex communication may have expanded. While nonhuman primates communicate in various ways, the growth of the human brain could have enabled the development of language, strengthening social bonds, and fostering increasingly complex cultural systems.<\/p>\n<p>In short, fossil evidence suggests that human evolution cannot be understood solely in biological terms. The emergence of bipedalism, brain expansion, and cultural behaviours are deeply interconnected, reflecting the intricate relationship between our physical evolution and the socio-cultural environments that shaped early hominins.<\/p>\n<h4>Discussion Questions<\/h4>\n<ul>\n<li>Based on the theories discussed above, what are your thoughts or opinions on the emergence of bipedalism? Has it benefited us in any way? What are some of the problems we face?<\/li>\n<li>What other cultural adaptations might have influenced our morphological features? Can we observe some of these changes today?<\/li>\n<li>Do you believe technology today is impacting our evolutionary journey? Please elaborate.<\/li>\n<\/ul>\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 metres tall and females at 1.24 metres.<\/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>Summary<\/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\"><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 centre of gravity (i.e., between the feet) in bipeds.<\/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>Aiello, L. C.; Wheeler, P. (1995).The Expensive-Tissue Hypothesis: The Brain and the Digestive System in Human and Primate Evolution. <em>Current Anthropology<\/em>, 36 (2):199-221<\/p>\n<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. 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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> 12 (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\">Van Le, Q., Isbell, L. A., Matsumoto, J., Nguyen, M., Hori, E., Maior, R. S., Tomaz, C., Tran, A. H., Ono, T., &amp; Nishijo, H. (2013). Pulvinar neurons reveal neurobiological evidence of past selection for rapid detection of snakes. <em>Proceedings of the National Academy of Sciences, 110<\/em>(47), 19000\u201319005. <\/span><\/p>\n<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>Wrangham, R.W. et al. (1999). \"The Raw and the Stolen\".\u00a0<em>Current Anthropology<\/em>.\u00a040\u00a0(5):\u00a0567\u2013594.\u00a0<a href=\"https:\/\/en.wikipedia.org\/wiki\/Doi_(identifier)\">doi<\/a>:<a href=\"https:\/\/doi.org\/10.1086%2F300083\">10.1086\/300083<\/a>.<\/p>\n<p>Wrangham, Richard. (2009). <em>Catching Fire: How Cooking Made Us Human.<\/em>\u00a0New York, NY: Basic Books.<\/p>\n<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>\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_1683_1688\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1688\"><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\">Learning Objectives<\/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 and 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-coloured 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 11.1). 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><img class=\"aligncenter\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image13-1.jpg\" alt=\"Reconstructed head and shoulders of a young Homo erectus.\" width=\"405\" height=\"308\" \/><figcaption class=\"wp-caption-text\">Figure 11.1a-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 10 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<\/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<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 8, 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 10, 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_1683_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_1683_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 11.2, 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 11.2: 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. It is speculated that 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 11.3) 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<figure style=\"width: 583px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 11.3: 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 undoubtedly challenging for our ancestors, but it offered new opportunities to make a living. It is theorized that 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 10) likely developed their large molar teeth with thick enamel in order to exploit this particular dietary niche.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Members of the genus <em>Homo <\/em>took a different route. Faced with the unstable African climate and shifting landscape, it is proposed that they evolved bigger brains that might have enabled them to rely on cultural solutions such as crafting stone tools that opened up new foraging opportunities. This strategy of behavioural flexibility might have served them well during this unpredictable time and may have 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 centimetres (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<figure style=\"width: 554px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image16.jpg\" alt=\"Africa map with South Africa, Tanzania, Kenya, and Ethiopia shaded.\" width=\"554\" height=\"717\" \/><figcaption class=\"wp-caption-text\">Figure 11.4: 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<p class=\"import-Normal\"><span style=\"color: #000000\">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 11.4). 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 analyse the remains. The fossil of the juvenile shown in Figure 11.5 (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<\/div>\n<figure id=\"attachment_316\" aria-describedby=\"caption-attachment-316\" style=\"width: 641px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-316\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/10.5.jpg\" alt=\"Two lateral right views of skulls and a jaw with blackened teeth.\" width=\"641\" height=\"214\" \/><figcaption id=\"caption-attachment-316\" class=\"wp-caption-text\">Figure 11.5a-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 the 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 11.6). 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 11.7). Researchers who favoured keeping all fossils in <em>Homo habilis<\/em> argued that sexual dimorphism, adaptation to local environments, or<strong> <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1566\">developmental plasticity<\/a><\/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 11.6: 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image10-1.jpg\" alt=\"Front view of black and white skull, missing lower jawbone.\" width=\"228\" height=\"228\" \/><figcaption class=\"wp-caption-text\">Figure 11.7: 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 11.8). 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 11.8: 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 11.5), 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\">It is theorized that 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. As discussed in Chapter 10, 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_1683_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 11.9). 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 11.9: 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 behaviour 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<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><\/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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image6-2.png\" alt=\"Lateral view of skull with large brow ridges.\" width=\"289\" height=\"289\" \/><figcaption class=\"wp-caption-text\">Figure 11.10: 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 11.10). 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_1683_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_1683_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_1683_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.<\/span><\/p>\n<div class=\"__UNKNOWN__\">\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 colour gene. Since chimpanzees have light skin under their hair, it is probable that early hominins also had light skin colour. 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<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 11.1). 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 11.11 and 11.12.<\/span><\/p>\n<figure style=\"width: 594px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 11.11:\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 11.12: 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 centre 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>\n<h2 class=\"__UNKNOWN__\"><span style=\"color: #000000\"><em>Homo erectus <\/em>Lifeways<\/span><\/h2>\n<div class=\"__UNKNOWN__\">\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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 11.13: 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_1683_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 11.13. 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<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Subsistence and Diet<\/strong><\/span><\/h3>\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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5-2.png\" alt=\"Five men excavating and note-taking at an archaeological site.\" width=\"253\" height=\"380\" \/><figcaption class=\"wp-caption-text\">Figure 11.14: 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 11.14), 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. 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, researchers propose that hominins began to cut up, tenderize, and process meat and plants, such that they did not have to use their teeth so vigorously.<\/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>It is proposed that the larger brain and body size of <em>Homo erectus<\/em> were fuelled 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. It is suggested that 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\u2014 might have given them greater flexibility in adapting to different environments. 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_1683_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_1683_1559\">human behavioural ecology<\/a><\/strong>, explored in more detail in Appendix C, examines the roots of human behaviour 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 11.15: 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 behavioural ecologists who study modern hunter-gatherer societies have observed that foraging is no easy business (Figure 11.15). 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_1683_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><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1560\">mosaic evolution<\/a><\/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 12), 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 behaviour 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 behavioural advancements, it now looks like many behaviours 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 behaviours; 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 behavioural 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 behaviours 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 behavioural ecology<\/strong>: The study of human behaviour 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<\/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>. Produced 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. 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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. 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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\">Acknowledgements<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_1683_1689\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1689\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<div class=\"textbox\">\n<h6>Student contributors to this chapter: Peyton Dagg, Bryana Henry, Anoriel Jacques, Saehee Yoon, and Brandon Hawke<\/h6>\n<p>Learn more about <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/about-the-authors\/\">authors and editors<\/a>,<a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/student-contributors\/\"> Hess' student contributions<\/a>, and <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/versioning-history\/\">versioning history<\/a>.<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\">Learning Objectives<\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li class=\"import-Normal\">Identify the main groupings of Archaic <em>Homo sapiens<\/em>, such as Neanderthals.<\/li>\n<li class=\"import-Normal\">Explain how shifting environmental conditions required flexibility of adaptations, both anatomically and behaviourally.<\/li>\n<li class=\"import-Normal\">Describe the unique anatomical and cultural characteristics of Archaic <em>Homo sapiens, <\/em>including Neanderthals, in contrast to other hominins.<\/li>\n<li class=\"import-Normal\">Articulate how Middle Pleistocene hominin fossils fit into evolutionary trends including cranial capacity (brain size) development, cultural innovations, and migration patterns.<\/li>\n<li class=\"import-Normal\">Identify the shared traits, regional variations, and local adaptations among Archaic <em>Homo sapiens.<\/em><\/li>\n<li class=\"import-Normal\">Detail the increased complexity and debates surrounding the classification of hominins in light of transitional species, species admixture, etc.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<h2 class=\"import-Normal\">Breaking the Stigma of the \"Caveman\"<\/h2>\n<p class=\"import-Normal\">What do you think of when you hear the word \u201ccaveman\u201d? Perhaps you imagine a character from a film such as <em>The Croods<\/em>, <em>Tarzan<\/em>, and <em>Encino Man<\/em> or from the cartoon <em>The Flintstones<\/em>. Maybe you picture the tennis-playing, therapy-going hairy Neanderthals from Geico Insurance commercials. Or perhaps you imagine characters from <em>The Far Side<\/em> or <em>B.C.<\/em> comics. Whichever you picture, the character in your mind is likely stooped over with a heavy brow, tangled long locks and other body hair, and clothed in animal skins, if anything. They might be holding a club with a confused look on their face, standing at the entrance to a cave or dragging an animal carcass to a fire for their next meal (see Figure 12.1). You might have even signed up to take this course because of what you knew\u2014or expected to learn\u2014about \u201ccavemen.\u201d<\/p>\n<figure style=\"width: 163px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/cdn.pixabay.com\/photo\/2013\/07\/13\/12\/36\/caveman-159964_1280.png\" alt=\"Free caveman beard man vector\" width=\"163\" height=\"317\" \/><figcaption class=\"wp-caption-text\">Figure 12.1: Cartoon illustration of a prehistoric \u201ccaveman\u201d, commonly used as a visual representation of early humans in the Stone Age. Credit: <em data-start=\"261\" data-end=\"293\">Caveman, Beard, Man, Primitive<\/em> free to use under the Pixabay <a href=\"https:\/\/pixabay.com\/service\/license-summary\/\">Content License.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\">These images have long been the stigma and expectation about our ancestors at the transition to modern <em>Homo sapiens<\/em>. Tracing back to works as early as Carl Linnaeus, scientists once propagated and advanced this imagery, creating a clear picture in the minds of early scholars that informed the general public, even through today, that Archaic <em>Homo sapiens<\/em>, \u201ccavemen,\u201d were somehow fundamentally different and much less intelligent than we are now. Unfortunately, this view is overly simplistic, misleading, and incorrect. Understanding what Archaic <em>Homo sapiens<\/em> were actually like requires a much more complex and nuanced picture, one that comes into sharper focus as continuing research uncovers more about the lives of our not-too-distant (and not-too-different) ancestors.<\/p>\n<p class=\"import-Normal\">The first characterizations of Archaic <em>Homo sapiens<\/em> were formed from limited fossil evidence in a time when <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1582\">ethnocentric<\/a><\/strong> and species-centric perspectives (<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1583\">anthropocentrism<\/a><\/strong>) were more widely accepted and entrenched in both society and science. Today, scientists are working from a more complete fossil record from three continents (Africa, Asia, and Europe), and genetic evidence informs their analyses and conclusions. The existence of Archaic <em>Homo sapiens<\/em> marks an exciting point in our lineage\u2014a point at which many modern traits had emerged and key refinements were on the horizon. Anatomically, humans today are not that much different from Archaic <em>Homo sapiens<\/em>.<\/p>\n<h2><\/h2>\n<h2 class=\"import-Normal\">The Changing Environment<\/h2>\n<p class=\"import-Normal\">While modern climate change is of critical concern today due to its cause (human activity) and pace (unprecedentedly rapid), the existence of global climate change itself is not a recent phenomenon. The focus of this chapter, the Middle Pleistocene (roughly between 780 kya and 125 kya), is the time period in which Archaic <em>Homo sapiens <\/em>appears in the fossil record\u2014a time that witnessed some of the most drastic climatic changes in human existence. During this time period, there were 15 major and 50 minor glacial events in Europe, alone.<\/p>\n<p class=\"import-Normal\">What exactly is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1635\">glaciation<\/a><\/strong>? When scientists talk about glacial events, they are referring to the climate being in an ice age. This means that the ocean levels were much lower than today, because much of the earth\u2019s water was tied up in large glaciers or ice sheets. Additionally, the average temperature would have been much cooler, which would have better supported an Arctic or tundra-adapted plant-and-animal ecosystem in northern latitudes. The most interesting and relevant features of Middle Pleistocene glacial events are the sheer number of them and their repeated bouts: this era alternated between glacial periods and warmer periods, known as<strong> interglacials<\/strong>. In other words, the planet wasn\u2019t in an ice age the whole time.<\/p>\n<p class=\"import-Normal\">You can see the dramatic and increasing fluctuations in temperature, recorded through <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_906\">foraminifera<\/a><\/strong>, in Figure 12.2. The distance between highs and lows demonstrates the severity of temperature shifts. Much as the Richter scale represents more intense earthquakes with more dramatic peaks, so too does this chart, which uses dramatic peaks to demonstrate intense temperature swings.<\/p>\n<figure id=\"attachment_346\" aria-describedby=\"caption-attachment-346\" style=\"width: 1753px\" class=\"wp-caption alignnone\"><img class=\"size-full wp-image-329\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/All_palaeotemps.png\" alt=\"The graph shows changes in Earth\u2019s temperature for the last 540 My.\" width=\"1753\" height=\"565\" \/><figcaption id=\"caption-attachment-346\" class=\"wp-caption-text\">Figure 12.2: The Geologic Timescale and corresponding temperature shifts. Wide and rapid shifts took place during the Pleistocene (the second box from the right). More dramatic fluctuations depict greater severity of temperature shift. The Eocene, Pliocene, and Holocene epochs had more stable temperatures. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:All_palaeotemps.png\">All paleotemps<\/a> by Glen Fergus 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\">Glacial periods are defined by Earth\u2019s average temperature being lower. Worldwide, temperatures are reduced, with cold areas becoming even colder. Huge portions of the landscape may have become inaccessible during glacial events due to the formation of glaciers and massive ice sheets. In Europe, the Scandinavian continental glacier covered what is today Ireland, England, Sweden, Norway, Denmark, and some of continental Europe. Plant and animal communities shifted to lower latitudes along the periphery of ice sheets. Additionally, some new land was opened during glacials. Evaporation with little runoff reduced sea levels by as much as almost 150 metres, shifting coastlines outward by in some instances as much as almost 100 kilometres. Additionally, land became exposed that connected what were previously unconnected continents such as Africa into Yemen at the Gulf of Aden.<\/p>\n<p class=\"import-Normal\">Glacial periods also affected equatorial regions and other regions that are today thought of as warmer or at least more temperate parts of the globe, including Africa. While these areas were not covered with glaciers, increased global glaciation resulted in lower sea levels and expanded coastlines. Cooler temperatures were accompanied by the drying of the climate, which caused significantly reduced rainfall, increased aridity, and the expansion of deserts. It is an interesting question to consider whether the same plants and animals that lived in these regions prior to the ice ages would be able to survive and thrive in this new climate. Plant and animal communities shifted in response to the changing climate, whenever possible.<\/p>\n<h2 class=\"import-Normal\">Surviving During the Middle Pleistocene<\/h2>\n<p class=\"import-Normal\">Rather than a single selective force, the Middle Pleistocene was marked by periods of fluctuation, not just cold periods. Interglacials interrupted glaciations, reversing trends in sea level, coastline, temperature, precipitation, and aridity, as well as glacier size and location. Interglacials are marked by increased rainfall and a higher temperature, which causes built-up ice in glaciers to melt. This leads to glacial retreat, which is the shrinking of glaciers and the movement of the glaciers back toward the poles, as we\u2019ve seen in our lifetime. During interglacials, sea levels increase, flooding some previously exposed coastlines and continental connections. In addition, plant and animal communities shift accordingly, often finding more temperate climates to the north and less arid and more humid climates in the tropics (Van Andel and Tzedakis 1996).<\/p>\n<p class=\"import-Normal\">Scientists have found that the Olorgesailie region in southern Kenya was at various times in the Middle Pleistocene a deep lake, a drought-dried lakebed with an area criss-crossed by small streams, and a grassland. While various animal species would have moved in and out of the area as the climate shifted, some animal species went extinct, and new, often related, species took up residence. The trend, scientists noted, was that animals with more specialized features went extinct and animals with more generalized features, such as animals we see today, survived in this changing climatic time period. For example, a zebra with specialized teeth for eating grass was ultimately replaced by a zebra that could eat both grass and other types of vegetation. If this small, localized example shows such a dramatic change in terms of the environment and the plant and animal biocommunities, what would have been the impact on humans?<\/p>\n<p class=\"import-Normal\">There is no way humans could have escaped the effects of Middle Pleistocene climate change, no matter what region of the world they were living in. As noted earlier, and as evidenced by what was seen in the other biotic communities, humans would have faced changing food sources as previous sources of food may have gone extinct or moved to a different latitude. Depending on where they were living, fresh water may have been limited. Durial glacials, lower sea levels would have given humans more land to live on, while the interglacials would have reduced the available land through the increase in rainfall and associated sea level rise. Dry land connections between the continents would have made movement from one continent to another by foot easier at times than today, although these passageways were not consistently available through the Middle Pleistocene due to the glacial\/interglacial cycle. Finally, as evidenced by the Olorgesailie region in Kenya, during the Middle Pleistocene animal species that were overly specialized to one particular type of environment were less likely to survive when compared to their more generalized counterparts. Evidence suggests that this same pattern may have held true for Archaic <em>Homo sapiens<\/em>, in terms of their ability to survive this dramatic period of climate change.<\/p>\n<h2 class=\"import-Normal\">Defining Characteristics of Archaic <em>Homo sapiens<\/em><\/h2>\n<p class=\"import-Normal\">Archaic <em>Homo sapiens<\/em> share our species name but are distinguished by the term \u201cArchaic\u201d as a way of recognizing both the long period of time between their appearance and ours, as well as the way in which human traits have continued to evolve over time\u2014making Archaic <em>Homo sapiens <\/em>look slightly different from us today, despite being considered the same species. Living throughout Africa, and the Middle East during the Middle Pleistocene, Archaic <em>Homo sapiens <\/em>are considered, in many ways, transitional between <em>Homo <\/em><em>erectus<\/em> and modern <em>Homo sapiens <\/em>(see Figure 12.3). Archaic<em> Homo sapiens<\/em> share the defining trait of an increased brain size of at least 1,100 cc and averaging 1,200 cc, although there are significant regional and temporal (time) variations. Because of these variations, scientists disagree on whether these fossils represent a single, variable species or multiple, closely related species (sometimes called <em>Homo antecessor<\/em>,<em> Homo bodoensis, Homo heidelbergensis<\/em>,<em> Homo georgicus<\/em>,<em> Homo neanderthalensis<\/em>, and <em>Homo rhodesiensis<\/em>).<\/p>\n<p class=\"import-Normal\">An active area of scholarship in the discipline involves reconciling the diversity of species from this time period and establishing the phylogenetic relationships between them. The term \u201cArchaic <em>Homo sapiens\u201d <\/em>can mean different things to different scholars within the discipline. The intent of this chapter is to provide an understanding of the diversity of this time period and provide data used to make interpretations from among the most likely possibilities. Although we recognize that some anthropologists split many of these fossils into separate species, until the issue is resolved at the discipline level, this chapter will rely on the widely used naming conventions that refer to many fossils from this time period as Archaic <em>Homo sapiens<\/em>. We will discuss these contemporaneous fossils as a unit<em>, <\/em>with the exception of a particularly unique population living in Europe and West Asia known as the Neanderthals, which we will examine separately.<\/p>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 468pt\">\n<caption>Figure 12.3: A comparison of Homo erectus, Archaic Homo sapiens, and anatomically modern Homo sapiens. This table compares key traits of the crania and postcrania that distinguish these three hominins. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-16\/\">Homo erectus, Archaic Homo sapiens, and anatomically modern Homo sapiens table (Figure 11.3)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Amanda Wolcott Paskey and AnnMarie Beasley Cisneros 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=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><strong>Trait<\/strong><\/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\"><strong><em>Homo <\/em><\/strong><strong><em>erectus<\/em><\/strong><\/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\"><strong>Archaic <\/strong><strong><em>Homo sapiens <\/em><\/strong><strong>(including Neaderthals)<\/strong><\/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\"><strong>Anatomically Modern <\/strong><strong><em>Homo sapiens<\/em><\/strong><\/p>\n<\/td>\n<\/tr>\n<\/thead>\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\"><strong>Time<\/strong><\/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\">1.8 mya\u2013200,000 ya<\/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\">600,000\u201340,000 ya<\/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\">315,000 ya\u2013today<\/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\"><strong>Brain size<\/strong><\/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\">900 cc<\/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\">1,200 cc (1,500 cc when including Neanderthals)<\/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\">1,400 cc<\/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\"><strong>Skull Shape<\/strong><\/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\">Long and low,<\/p>\n<p class=\"import-Normal\">angular<\/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\">Intermediate<\/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\">Short and high, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1643\">globular<\/a><\/strong><\/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\"><strong>Forehead<\/strong><\/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\">Absent<\/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\">Emerging<\/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\">Present<\/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\"><strong>Nasal Region<\/strong><\/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\">Projecting nasal bones (bridge of the nose), no midfacial prognathism<\/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\">Wider nasal aperture and some midfacial prognathism, particularly pronounced among Neanderthals<\/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\">Narrower nasal aperture, no midfacial prognathism<\/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\"><strong>Chin<\/strong><\/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\">Absent<\/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\">Absent<\/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\">Present<\/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\"><strong>Other Facial Features<\/strong><\/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\">Large brow ridge and large projecting face<\/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\">Intermediate<\/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\">Small brow ridge and<strong> retracted face<\/strong><\/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\"><strong>Other Skull Features<\/strong><\/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\">Nuchal torus, sagittal keel, thick cranial bone<\/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\">Projecting occipital bone, often called occipital bun in Neanderthals; intermediate thickness of cranial bone<\/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\">Small bump on rear of skull, if anything; thin cranial bone<\/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\"><strong>Dentition<\/strong><\/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\">Large teeth, especially front teeth<\/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\">Slightly smaller teeth; front teeth still large;<\/p>\n<p class=\"import-Normal\">retromolar gap in Neanderthals<\/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\">Smaller teeth<\/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\"><strong>Postcranial Features<\/strong><\/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\">Robust bones of skeleton<\/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\">Robust bones of skeleton<\/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\">More gracile bones of skeleton<\/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<p class=\"import-Normal\">When comparing <em>Homo <\/em><em>erectus<\/em>, Archaic<em> Homo sapiens, <\/em>and anatomically modern <em>Homo sapiens<\/em>, one can see that Archaic<em> Homo sapiens<\/em> are intermediate in their physical form. For some features, this follows the trends first seen in <em>Homo <\/em><em>erectus<\/em> with other features having early, less developed forms of traits seen in modern <em>Homo sapiens<\/em>. For example, Archaic <em>Homo sapiens<\/em> trended toward less angular and higher skulls than <em>Homo <\/em><em>erectus<\/em><em>. <\/em>However, the archaic skulls were not as short and globular and had less developed foreheads compared to anatomically modern <em>Homo sapiens. <\/em>Archaic <em>Homo sapiens<\/em> had smaller brow ridges and a less-projecting face than <em>Homo <\/em><em>erectus<\/em> and slightly smaller teeth, although incisors and canines were often about as large as those of <em>Homo <\/em><em>erectus<\/em>. Archaic <em>Homo sapiens <\/em>also had a wider <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1638\">nasal aperture<\/a><\/strong>, or opening for the nose, and a forward-projecting midfacial region, which is later seen more fully developed among Neanderthals and is known as <strong>midfacial prognathism<\/strong>. The occipital bone often projected and the cranial bone was of intermediate thickness, somewhat reduced from <em>Homo <\/em><em>erectus<\/em> but not nearly as thin as that of anatomically modern <em>Homo sapiens. <\/em>The postcrania remained fairly robust. Identifying a set of features that is unique to Archaic<em> Homo sapiens<\/em> is a challenging task, due to both individual and geographic variation\u2014these developments were not all present to the same degree in all individuals. Neanderthals are the exception, as they had several unique traits that made them notably different from modern <em>Homo sapiens<\/em> as well as their closely related Archaic cousins.<\/p>\n<figure style=\"width: 299px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image10-4.png\" alt=\"Archaic Homo sapiens skull cast.\" width=\"299\" height=\"299\" \/><figcaption class=\"wp-caption-text\">Figure 12.4: \u201cBroken Hill Man,\u201d found at Kabwe in Zambia, shows common traits associated with archaic Homo sapiens in Africa, including a large brain, taller cranium, and Homo erectus-like features such as massive brow ridges, a large face, and thick cranial bones. Credit: <a href=\"https:\/\/boneclones.com\/product\/homo-heidelbergensis-skull-broken-hill-1-rhodesian-man-BH-004\">Homo heidelbergensis Cranium Broken Hill 1 (Rhodesian Man)<\/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 one thing that is clear about Archaic <em>Homo sapiens<\/em> is that, despite general features, there is a lot of regional variation, which is first seen in the different <em>Homo <\/em><em>erectus<\/em> specimens across Asia and Africa. While the general features of Archaic <em>Homo sapiens<\/em>, identified earlier, are present in the fossils of this time period, there are significant regional differences. The majority of this regional variation lies in the degree to which fossils have features more closely aligned with <em>Homo <\/em><em>erectus<\/em> or with anatomically modern <em>Homo sapiens<\/em>.<\/p>\n<figure style=\"width: 244px\" class=\"wp-caption alignright\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image7-5.png\" alt=\"Side view of the Dali cranium.\" width=\"244\" height=\"213\" \/><figcaption class=\"wp-caption-text\">Figure 12.5: Dali cranium, found at Dali, China, is representative of traits seen in archaic Homo sapiens in Asia, including large and robust features with heavy brow ridges like Homo erectus and a large cranial capacity intermediate between Homo erectus and anatomically modern Homo sapiens. Credit: Dali skull original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by <a href=\"https:\/\/marynelsonstudio.com\">Mary Nelson<\/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>To illustrate this point, we will examine three exemplary specimens, one from each of the three continents on which Archaic<em> Homo sapiens <\/em>lived. First, in Africa, a specimen from Broken Hill is one of several individuals found in the Kabwe lead mine in Zambia. It had a large brain (1,300 cc) and taller cranium as well as many <em>Homo erectus-<\/em>like skull features, including massive brow ridges, a large face, and thick cranial bones (Figure 12.4). Second, one partial crania from Dali, China, is representative of Archaic<em> Homo sapiens <\/em>in Asia, with large and robust features with heavy brow ridges, akin to what is seen in <em>Homo <\/em><em>erectus<\/em>, and a large cranial capacity intermediate between <em>Homo <\/em><em>erectus<\/em> and anatomically modern <em>Homo sapiens<\/em> (Figure 12.5). Third, an almost-complete skeleton was found in northern Spain at Atapuerca. Atapuerca 5 (Figure 12.6) has thick cranial bone, an enlarged cranial capacity, intermediate cranial height, and a more rounded cranium than seen previously. Additionally, Atapuerca 5 demonstrates features that foreshadow Neanderthals, including increased midfacial prognathism. After examining some of the fossils such as those from Kabwe, Dali, and Atapuerca, the transitional nature of Archaic<em> Homo sapiens <\/em>is clear: their features place them squarely between <em>Homo <\/em><em>erectus<\/em> and modern <em>Homo sapiens<\/em>.<\/p>\n<figure style=\"width: 293px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image13-4.png\" alt=\"Archaic Homo sapiens skull cast with mandible.\" width=\"293\" height=\"293\" \/><figcaption class=\"wp-caption-text\">Figure 12.6: Atapuerca 5 archaic Homo sapiens, found in northern Spain, is representative of traits seen in archaic Homo sapiens in Europe, including a thick cranial bone, an enlarged cranial capacity, intermediate cranial height, a more rounded cranium, and increased midfacial projection. Credit: <a href=\"https:\/\/boneclones.com\/product\/homo-heidelbergensis-skull-atapuerca-5-BH-022\">Homo heidelbergensis Skull Atapuerca 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 class=\"import-Normal\">Due to the transitional nature of Archaic<em> Homo sapiens<\/em>, identifying the time period with which they are associated is problematic and complex. Generally, it is agreed that Archaic<em> Homo sapiens <\/em>lived between 600,000 and 200,000 years ago, with regional variation and overlap between <em>Homo <\/em><em>erectus<\/em> on the early end of the spectrum and modern <em>Homo sapiens <\/em>and Neanderthals on the latter end. The earliest-known Archaic<em> Homo sapiens <\/em>fossils tentatively date to about 600,000 years ago in Africa, to around 300,000 years ago in Asia, and to about 350,000 years ago in Europe (and potentially as early as 600,000 years ago). Determining the end point of Archaic<em> Homo sapiens <\/em>is also problematic since it largely depends upon when the next subspecies of <em>Homo sapiens <\/em>appears and the classification of highly intermediate specimens. For example, in Africa, the end of Archaic<em> Homo sapiens <\/em>is met with the appearance of modern <em>Homo sapiens<\/em>, while in Europe it is the appearance of Neanderthals that is traditionally seen as marking the transition from other Archaic<em> Homo sapiens. <\/em><\/p>\n<p class=\"import-Normal\">It is important to remember that this time period is represented by many branching relationships and assuming an evolutionary trajectory that follows a single linear path would not be correct. Even still, Archaic<em> Homo sapiens <\/em>mark an important chapter in the human lineage, connecting more ancestral forms, such as <em>Homo <\/em><em>erectus<\/em>, to modern <em>Homo sapiens<\/em>. During this period of climatic transition and fluctuation, Archaic <em>Homo sapiens<\/em> mirror the challenges of their environments. Showing increasing regional variation due to the need for local adaptation, there is no single archetype for this group; the defining characteristic seems to be variability.<\/p>\n<h2 class=\"import-Normal\">Neanderthals<\/h2>\n<p class=\"import-Normal\">One well-known population of Archaic <em>Homo sapiens <\/em>are the Neanderthals, named after the site where they were first discovered in the Neander Valley, or \u201cthal\u201d in German, located near Dusseldorf, Germany. Popularly known as the stereotypical \u201ccavemen\u201d examined at the outset of this chapter, recent research is upending long-held beliefs about this group of Archaics. Neanderthal behaviour was increasingly complex, far beyond what was exhibited by even other Archaic <em>Homo sapiens<\/em> discussed throughout this chapter. We implore you to forget the image of the iconic caveman and have an open mind when exploring the fossil evidence of the Neanderthals.<\/p>\n<p class=\"import-Normal\">It is important to understand why Neanderthals are separated from other Archaic <em>Homo sapiens<\/em>. Unlike the rest of Archaic <em>Homo sapiens<\/em>, Neanderthals are easily defined and identified in many ways. Evidence suggests the time period when Neanderthals lived was between 150,000 and 40,000 years ago. There is a clear geographic boundary of where Neanderthals lived: western Europe, the Middle East, and western Asia. No Neanderthal fossils have ever been discovered outside of this area, including Africa. This is a bit curious, as other Archaics seem to have adapted in Africa and then migrated elsewhere, but Neanderthals\u2019 regional association makes sense in light of the environment to which they were best adapted: namely, extreme cold weather. Additionally, Neanderthals have a unique and distinct cluster of physical characteristics. While a few aspects of Neanderthals are shared among some Archaic <em>Homo sapiens<\/em>, such as the types of tools, most Neanderthal anatomical and behavioural attributes are unique to them.<\/p>\n<p class=\"import-Normal\">Neanderthals lived during some of the coldest times during the last Ice Age and at far northern latitudes. This means Neanderthals were living very close to the glacial edge, rather than in a more temperate region of the globe like some of their Archaic <em>Homo sapiens<\/em> relatives. While able to survive in arctic conditions, most Neanderthal sites dating to the glacial periods were found farther away from the severe cold, in a steppe tundra-like environment, which would have been more hospitable to Neanderthals, and their food sources, both flora and fauna (Ashton 2002; Nicholson 2017; Richter 2006). Their range likely expanded and contracted along with European glacial events, moving into the Middle East during glacial events when Europe became even cooler, and when the animals they hunted would have moved for the same reason. During interglacials, when Europe warmed a bit, Neanderthals and their prey would have been able to move back into Western Europe. Clearly, the true hallmark of Neanderthals is their adaptation to an unstable environment, shifting between warm and cold, as the climate was in constant flux throughout their existence (Adler Et al. 2003; Boettger Et al. 2009).<\/p>\n<p class=\"import-Normal\">Many of the Neanderthals\u2019 defining physical features are more extreme and robust versions of traits seen in other Archaic<em> Homo sapiens<\/em>, clustered in this single population. Brain size, namely an enlargement of the cranial capacity, is one such trait. The average Neanderthal brain size is around 1,500 cc, and the range for Neanderthal brains can extend to upwards of 1,700 cc. The majority of the increase in the brain occurs in the occipital region, or the back part of the brain, resulting in a skull that has a large cranial capacity with a distinctly long and low shape that is slightly wider than previous forms at the far back of the skull. Modern humans have a brain size comparable to that of Neanderthals; however, our brain expansion occurred in the frontal region of the brain, not the back, as in Neanderthal brains. This difference is also the main reason why Neanderthals lack the vertical forehead that modern humans possess. They simply did not need an enlarged forehead, because their brain expansion occurred in the rear of their brain. Due to cranial expansion, the back of the Neanderthal skull is less angular (as compared to <em>Homo erectus<\/em>), but not as rounded as <em>Homo sapiens<\/em>, producing an elongated shape, akin to a football.<\/p>\n<p class=\"import-Normal\">Another feature that continues the trend noted in previous hominins is the enlargement of the nasal region, or the nose. Neanderthal noses are large and have a wide nasal aperture, which is the opening for the nose. While the nose is only made up of two bones, the nasals, the true size of the nose can be determined by looking at other facial features, including the nasal aperture, and the angle of the nasal and maxillary, or facial bones. In Neanderthals, these indicate a large, forward-projecting nose that appears to be pulled forward away from the rest of the face. This feature is further emphasized by the backward-sloping nature of the cheekbones, or the zygomatic arches. The unique shape and size of the Neanderthal nose is often characterized by the term <em>midfacial prognathism<\/em>\u2014a jutting out of the middle portion of the face, or nose. This is in sharp contrast to the prognathism exhibited by other hominins, who exhibited prognathism, or the jutting out, of their jaws.<\/p>\n<p class=\"import-Normal\">The teeth of the Neanderthals follow a similar pattern seen in the Archaic <em>Homo sapiens<\/em>, which is an overall reduction in size, especially as compared to the extremely large teeth seen in the genus <em>Australopithecus<\/em>. However, while the teeth continued to reduce, the jaw size did not keep pace, leaving Neanderthals with an oversized jaw for their teeth, and a gap between their final molar and the end of their jaw. This gap is called a <strong>retromolar gap<\/strong>.<\/p>\n<p class=\"import-Normal\">The projecting occipital bone present in other Archaic<em> Homo sapiens <\/em>is also more prominent in Neanderthals, extending the trend found in Archaics. Among Neanderthals, this projection of bone is easily identified by its bun shape on the back of the skull and is known as an <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1645\">occipital bun<\/a><\/strong>. This projection appears quite similar to a dinner roll in size and shape. Its purpose, if any, remains unknown.<\/p>\n<p class=\"import-Normal\">Continuing the Archaic<em> Homo sapiens <\/em>trend, Neanderthal brow ridges are prominent but somewhat smaller in size than those of <em>Homo <\/em><em>erectus<\/em> and earlier Archaic<em> Homo sapiens. <\/em>In Neanderthals, the brow ridges are also often slightly less arched than those of other Archaic<em> Homo sapiens<\/em>.<\/p>\n<p class=\"import-Normal\">In addition to extending traits present in Archaic<em> Homo sapiens, <\/em>Neanderthals possess several distinct traits. Neanderthal <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1646\">infraorbital foramina<\/a><\/strong>, the holes in the maxillae or cheek bones through which blood vessels pass, are notably enlarged compared to other hominins. The Neanderthal postcrania are also unique in that they demonstrate increased robusticity in terms of the thickness of bones and body proportions that show a barrel-shaped chest and short, stocky limbs, as well as increased musculature. These body portions are seen across the spectrum of Neanderthals\u2014in men, women, and children.<\/p>\n<p class=\"import-Normal\">Traditionally, many of the unique traits that Neanderthals possess were seen as adaptations to the extreme cold, dry environments in which they often lived and which exerted strong selective forces. For example, Bergmann\u2019s and Allen\u2019s Rules dictate that an increased body mass and short, stocky limbs are common in animals that live in cold conditions. Neanderthals were said to have matched the predictions of Bergmann\u2019s and Allen\u2019s Rules perfectly (Churchill 2006). In addition, the Neanderthal skull also exhibits adaptations to the cold. Neanderthals\u2019 large infraorbital foramina allow for larger blood vessels, increasing the volume of blood that is found closest to the skin, which helps to keep the skin warmer. Their enlarged noses resulted in longer nasal passages and mucus membranes that warmed and moistened cold air before it reached the lungs. The Neanderthals\u2019 larger nose has long been thought to have acted as a humidifier, easing physical exertion in their climate, although research on this particular trait continues to be studied and debated (Rae Et al. 2011).<\/p>\n<p class=\"import-Normal\">New research, however, seems to suggest that these unique skeletal adaptations might not have been strict adaptations to cold weather (Evteev Et al. 2017; Pearce Et al. 2013). For example, large brow ridges might have served as a way to shade the face from the sun. The increased occipital portion of the brain, some researchers state, was to support a larger visual system present in Neanderthals. This visual system would have given them increased light sensitivity, which would have been useful in higher latitudes that had dark winters. And, while recent modeling of nostril airflow on reconstructed Neanderthal specimens supports the notion that Neanderthals had extensive mucus membranes inside their noses, the data shows that modern <em>Homo sapiens<\/em> are superior to Neanderthals in our ability to use our noses as a way to warm and cool air. However, Neanderthals were able to snort air through their noses better than we can. Why is this important? When combined with the fact that Neanderthals tended to prefer a more temperate, tundra-like environment, and that other physical traits suggest that Neanderthals had huge bodies that needed massive amounts of calories to sustain them, the picture gets clearer. Massive amounts of energy would have been required to power a Neanderthal body, and anything that might have made them more calorically efficient would have been favoured. Efficient breathing, through larger noses into large lungs, meaning deeper breaths, would have been favoured. To further save energy expenditure, body sizes might have been sacrificed as well. These same types of adaptations are similar to ones seen in children today who are born in high altitudes, not cold climates. Figure 12.7 provides a summary of these unique features of the Neanderthal.<\/p>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 468pt\">\n<caption>Figure 12.7: Neanderthal distinguishing features. This table outlines key features associated with Neanderthals. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-16\/\">Neanderthal distinguishing features table (Figure 11.6)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Amanda Wolcott Paskey and AnnMarie Beasley Cisneros 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: 21pt\">\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\" colspan=\"2\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Distinct Neanderthal Anatomical Features<\/strong><\/p>\n<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table2-R\" style=\"height: 0\">\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><strong>Brain Size<\/strong><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">1,500 cc average<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 0\">\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><strong>Skull Shape<\/strong><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Long and low<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 0\">\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><strong>Brow Ridge Size<\/strong><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Large<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 0\">\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><strong>Nose Size<\/strong><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Large, with midfacial prognathism<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 0\">\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><strong>Dentition<\/strong><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Reduced, but large jaw size, creating retromolar gap<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 0\">\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><strong>Occipital Region<\/strong><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Enlarged occipital region, occipital bun<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 0\">\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><strong>Other Unique Cranial Features<\/strong><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Large infraorbital foramina<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 0\">\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><strong>Postcranial Features<\/strong><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Short and stocky body, increased musculature, barrel-shaped chest<\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<p class=\"import-Normal\">A classic example of a Neanderthal with all of the characteristics mentioned above is the nearly complete La Ferrassie 1 Neanderthal, from France. This is a male skeleton, with a brain size of around 1640cc, an extremely large nose and infraorbital foramina, brow ridges that are marked in size, and an overall robust skeleton (Figure 12.8).<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 390px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image1-4.png\" alt=\"A reproduction of a complete Neanderthal skeleton.\" width=\"390\" height=\"689\" \/><figcaption class=\"wp-caption-text\">Figure 12.8: La Ferrassie 1 Neanderthal is representative of many classic Neanderthal features, including a large brain, large nose, large infraorbital foramina, large brow ridges, and robust postcrania. Credit: <a href=\"https:\/\/boneclones.com\/product\/neanderthal-skeleton-articulated-SC-019-A\">Neanderthal Skeleton Articulated<\/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<h3 class=\"import-Normal\"><strong>Neanderthal Culture: Tool Making and Use<\/strong><\/h3>\n<p class=\"import-Normal\">One key Neanderthal adaptation was their cultural innovations, which are an important way that hominins adapt to their environment. As you recall, <em>Homo erectus<\/em>'s tools, Acheulean handaxes, represented an increase in complexity over Oldowan tools, allowing more efficient removal of meat and possibly calculated scavenging. In contrast, Neanderthal tools mark a significant innovation in tool-making technique and use with <strong>Mousterian tools<\/strong> (named after the Le Moustier site in southwest France). These tools were significantly smaller, thinner, and lighter than Acheulean handaxes and formed a true toolkit. The materials used for Mousterian tools were of higher quality, which allowed for both more precise toolmaking and tool reworking when the tools broke or dulled after frequent reuse. The use of higher-quality materials is also indicative of required forethought and planning to acquire them for tool manufacture. It has been suggested that the Neanderthals, unlike <em>Homo <\/em><em>erectus<\/em>, saved and reused their tools, rather than making new ones each time a tool was needed.<\/p>\n<figure style=\"width: 290px\" class=\"wp-caption alignright\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image3-4.png\" alt=\"Large flakes separated from the core.\" width=\"290\" height=\"159\" \/><figcaption class=\"wp-caption-text\">Figure 12.9: The Levallois technique is used to create Mousterian tools. The multistep process involves preparing the core in a specific way to yield flakes that can be used as tools. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Nucl%C3%A9us_Levallois_La-Parrilla.png\">Nucl\u00e9us Levallois La-Parrilla<\/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<p>Mousterian tools are constructed in a very unique manner, utilizing the <strong>Levallois technique<\/strong> (Figure 12.9), named after the first finds of tools made with this technique, which were discovered in the Levallois-Perret suburb of Paris, France. The Levallois technique is a multistep process that requires preparing the core, or raw material, in a specific way that will yield flakes that are roughly uniform in dimension. The flakes are then turned into individual tools. The preparation of the core is akin to peeling a potato or carrot with a vegetable peeler\u2014when peeling vegetables, you want to remove the skin in long, regular strokes, so that you are taking off the same amount of the vegetable all the way around. In the same way, the Levallois technique requires removing all edges of the <strong>cortex<\/strong>, or outside surface of the raw material, in a circle before removing the lid. The flakes, which will eventually be turned into the individual tools, can then be removed from the core. The potential yield of tools from one core would be many, as seen in Figure 12.10, compared to all previous tool-making processes, in which one core yielded a single tool. This manufacturing process might be considered the ultimate zero-waste tool-making technique (Delpiano Et al. 2018).<\/p>\n<figure style=\"width: 589px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image19-4.png\" alt=\"Levallois core and flakes that are gray in color and various shapes and sizes.\" width=\"589\" height=\"470\" \/><figcaption class=\"wp-caption-text\">Figure 12.10: Levallois core and flakes for tool production. Using this technique, one core is used to produce many flakes, each of which can be turned into a tool. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:NHM_-_Levalloiskern.jpg\">NHM - Levalloiskern<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Xenophon\">Wolfgang Sauber<\/a> (user: Xenophon) 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\">It is suggested that Neanderthal tools were used for a variety of purposes, including cutting, butchering, woodworking or antler working, and hide working. Additionally, because the Mousterian tools were lighter than previous stone tools, Neanderthals could <strong>haft<\/strong>, or attach the tool onto a handle, as the stone would not have been too heavy (Degano Et al. 2019). Neanderthals attached small stone blades onto short wood or antler handles to make knives or other small weapons, as well as attached larger blades onto longer shafts to make spears. New research examining tar-covered stones and black lumps at several Neanderthal sites in Europe suggests that Neanderthals may have been making tar by distilling it from birch tree bark, which could have been used to glue the stone tool onto its handle. If Neanderthals were, in fact, manufacturing tar to act as glue, this would predate modern humans in Africa using tree resin or similar adhesives by nearly 100,000 years.<\/p>\n<p class=\"import-Normal\">Evidence shows that raw materials used by Neanderthals came from distances as far away as 100 km. This could indicate a variety of things regarding Neanderthal behaviour, including a limited trade network with other Neanderthal groups or simply a large area scoured by Neanderthals when collecting raw materials. While research on specific applications continues, it should be clear from this brief discussion that Neanderthal tool manufacturing was much more complex than previous tool-making efforts, requiring technical expertise, patience, and skills beyond toolmaking to carry out.<\/p>\n<h3 class=\"import-Normal\"><strong>Neanderthal Culture: Hunting and Diet<\/strong><\/h3>\n<p class=\"import-Normal\">With their more sophisticated suite of tools and robust muscular bodies, Neanderthals were better armed for hunting than previous hominins. The animal remains in Neanderthal sites show that, unlike earlier Archaic <em>Homo sapiens<\/em>, Neanderthals were very effective hunters who were able to kill their own prey, rather than relying on scavenging. Though more refined than the tools of earlier hominins, the Neanderthal spear was not the kind of weapon that would have been thrown; rather, it would have been used in a jabbing fashion (Churchill 1998; Kortlandt 2002). This may have required Neanderthals to hunt in groups rather than individually and made it necessary to approach their prey quite closely (Gaudzinski-Windheuser Et al. 2018). Remember, the animals living with Neanderthals were very large-bodied due to their adaptations to cold weather; this would have included species of deer, horses, and bovids (relatives of the cow).<\/p>\n<p class=\"import-Normal\">Isotopes from Neanderthal bones show that meat was a significant component of their diet, similar to that seen in carnivores like wolves (Bocherens Et al. 1999; Jaouen Et al. 2019; Richards Et al. 2000). In addition to large prey, their diet included ibex, seals, rabbits, and pigeons. Though red meat was a critical component of the Neanderthal diet, evidence shows that at times they also ate limpets, mussels, and pine nuts. Tartar examined from Neanderthal teeth in Iraq and Belgium reveal that they also ate plant material including wheat, barley, date palms, and tubers, first cooking them to make them palatable (Henry Et al. 2010). While Neanderthals\u2019 diet varied according to the specific environment in which they lived, meat comprised up to 80% of their diet (Wi\u1e9ein Et al. 2015).<\/p>\n<h3 class=\"import-Normal\"><strong>Neanderthal Culture: Caring for the Injured and Sick<\/strong><\/h3>\n<p class=\"import-Normal\">While the close-range style of hunting used by Neanderthals was effective, it also had some major consequences. Many Neanderthal skeletons have been found with significant injuries, which could have caused paralysis or severely limited their mobility. Many of the injuries are to the head, neck, or upper body. Thomas Berger and Erik Trinkaus (1995) conducted a statistical comparative analysis of Neanderthal injuries compared to those recorded in modern-day workers\u2019 compensation reports and found that the closest match was between Neanderthal injuries and those of rodeo workers. Rodeo professionals have a high rate of head and neck injuries that are similar to the Neanderthals\u2019 injuries. What do Neanderthals and rodeo workers have in common? They were both getting very close to large, strong animals, and at times their encounters went awry.<\/p>\n<p class=\"import-Normal\">The extensive injuries sustained by Neanderthals are evident in many fossil remains. Shanidar 1 (Figure 12.11), an adult male found at the Shanidar site in northern Iraq and dating to 45,000 ya, has a lifetime of injuries recorded in his bones (Stewart 1977). Shanidar 1 sustained\u2014and healed from\u2014an injury to the face that would have likely caused blindness. His lower right arm was missing and the right humerus shows severe atrophy, likely due to disuse. This pattern has been interpreted to indicate a substantial injury that required or otherwise resulted in amputation or wasting away of the lower arm. Additionally, Shanidar 1 suffered from bony growths in the inner ear that would have significantly impaired his hearing and severe arthritis in the feet. He also exhibited extensive anterior tooth wear, matching the pattern of wear found among modern populations who use their teeth as a tool. Rather than an anomaly, the type of injuries evident in Shanidar 1 are similar to those found in many other Neanderthal fossils, revealing injuries likely sustained from hunting large mammals as well as demonstrating a long life of physical activity.<\/p>\n<p>&nbsp;<\/p>\n<p><img class=\"aligncenter\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image11-4.png\" alt=\"Neaderthal skull.\" width=\"329\" height=\"329\" \/><\/p>\n<figure style=\"width: 330px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5-4.png\" alt=\"Neaderthal right and left humerus. The right humerus is withered looking.\" width=\"330\" height=\"330\" \/><figcaption class=\"wp-caption-text\">Figure 12.11a-b: a. The Shanidar 1 skull shows an injury to the face that would likely have caused blindness. b. The Shanidar 1 right humerus (on the left side of the image) shows severe atrophy, likely due to disuse. Credit: a. <a href=\"https:\/\/boneclones.com\/product\/shanidar-1-skull-BH-050\">Homo neanderthalensis Shanidar 1 Skull<\/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>. b. <a href=\"https:\/\/humanorigins.si.edu\/evidence\/human-fossils\/fossils\/shanidar-1\">Shanidar 1<\/a> by Chip Clark, <a href=\"https:\/\/www.si.edu\/\">Smithsonian Institution<\/a> [exhibit: <a href=\"https:\/\/humanorigins.si.edu\/research\">Human Evolution <\/a>Evidence, Human Fossils, Species, Homo neanderthalensis] <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\">The pattern of injuries is as significant as the fact that Shanidar 1 and other injured Neanderthals show evidence of having <em>survived<\/em> their severe injuries. One of the earliest-known Neanderthal discoveries\u2014the one on whom misinformed analysis shaped the stereotype of the species for nearly a century\u2014is the La Chapelle-aux-Saints Neanderthal (Trinkaus 1985). The La Chapelle Neanderthal had a damaged eye orbit that likely caused blindness and suffered arthritis of the spine (Dawson and Trinkaus 1997). He had also lost most of his teeth, many of which he had lived without for so long that the mandibular and maxillary bones were partially reabsorbed due to lack of use. The La Chapelle Neanderthal was also thought to be at least in his mid-forties at death, an old age for the rough life of the Late Pleistocene\u2014giving rise to his nickname, \u201cthe Old Man.\u201d To have survived so long with so many injuries that obviously precluded successful large game hunting, he must have been taken care of by others. Such caretaking behaviour is also evident in the survival of other seriously injured Neanderthals, such as Shanidar 1. Long thought to be a hallmark modern human characteristic, taking care of the injured and elderly, for example preparing or pre-chewing food for those without teeth, indicates strong social ties among Neanderthals.<\/p>\n<h3 class=\"import-Normal\"><strong>Neanderthal Culture: Ritual Life<\/strong><\/h3>\n<p class=\"import-Normal\">Such care practices may have been expressed upon death as well. Nearly complete Neanderthal skeletons are not uncommon in the fossil record, and most are well preserved within apparently deliberate burials that involve deep graves and bodies found in specific, often fetal or <strong>flexed positions<\/strong> (Harrold 1980)<strong>.<\/strong> Discoveries of pollen in a grave at the Shanidar site in the 1960s led scientists to think that perhaps Neanderthals had placed flowering plants in the grave, an indication of ritual ceremony or spirituality so common in modern humans. But more recent investigations have raised some doubt about this conclusion (Pomeroy Et al. 2020). The pollen may have been brought in by burrowing rodents. Claims of <strong>grave goods<\/strong> or other ornamentation in burials are similarly debated, although possible.<\/p>\n<p class=\"import-Normal\">Some tantalizing evidence for symbolism, and debatably, ritual, is the frequent occurrence of natural pigments, such as <strong>ochre<\/strong> (red) and manganese dioxide (black) in Neanderthal sites that could have been used for art. However, the actual uses of pigments are unclear, as there is very little evidence of art or paintings in Mousterian sites. One exception may be the recent discovery in Spain of a perforated shell that appears to be painted with an orange pigment, which may be evidence of Neanderthal art and jewelry. However, many pigments also have properties that make them good emulsifiers in adhesive (like for attaching a stone tool to a wooden handle) or useful in tanning hides. So the presence of pigment may or may not be associated with symbolic thought; however, it definitely does show a technological sophistication beyond that of earlier Archaic hominins.<\/p>\n<div class=\"textbox shaded\" style=\"background: var(--lightblue)\">\n<h2>Dig Deeper: Evidence of Endocannibalism Among the Neanderthals<\/h2>\n<p>Krapina, a Neanderthal site in Croatia, has recently sparked new archeological discourse as many investigations upon fossilized remains show evidence of post-mortem modifications and manipulations of limbs and bones through the use of tools (Rougier Et al., 2016). These findings provide compelling evidence of Neanderthals engaging in cannibalism as part of their post-mortem practices. Additionally, these uncovered remains were found to have died of natural causes as opposed to being killed, showing that even the earliest humans may have had some sort of morality or ethics surrounding the cannibalizing of their kin, meaning they were aware of death in a social context as opposed to merely a physical one.<\/p>\n<p>While the original evidence in Krapina was uncovered in 1901, Croatian geologist and paleontologist Dragutin Gorjanovi\u0107-Kramberger\u2019s discovery of fragmented and burned human bones (Ullrich, 2005) was not yet confirmed to be linked to endocannibalism until much later. Whereas the discovery of burned bones does not mean they were being prepared for consummation, due to its context among other findings, this information supports the hypothesis that early hominids conducted post-mortem rituals and practices with their dead. Building on Gorjanovi\u0107-Kramberger\u2019s research, Herbert Ullrich wrote in <em>Anthropologie<\/em> (2005) that broken bones\u2014resulting from post-mortem bodily manipulations\u2014were \u201cdefleshed in preparation for secondary burial\u201d (2005, 251) and intentionally left outside rock shelters, while selectively chosen bones were seemingly brought inside for use in mortuary practices.<\/p>\n<p>For nearly 150 years, since the first Neanderthal skeletal remains were discovered, anthropologists and researchers have continued to debate the cognitive, social, and physical abilities of this species. In 2016, Rougier and colleagues wrote in <em>Scientific Reports<\/em>, furthering the research, presenting 99 new Neanderthal remains found in Goyet, Belgium. Among these remains, similar evidence of human-induced alterations was identified, including signs of butchering, consumption, and the use of bones to modify stone tools (Rougier Et al., 2016). This discovery provides significant support for the presence of cannibalistic behaviour among Northern European Neanderthals, contributing to the growing body of evidence that Neanderthals engaged with death in ways that reflect social awareness, ritual behaviour, and complex cultural practices.<\/p>\n<p><strong>Contemporary Cases of Prion Disease Related to Endocannibalism<\/strong><\/p>\n<p>The evidence of endocannibalism does not end with early hominids; with Australian medical anthropologists recording thousands of cannibalism-related prion disease occurrences present in populations up until 2009 (Radford &amp; Scragg, 2013). Following a mysterious epidemic of a new form of spongiform encephalopathy\u2013a \u201cprogressive degenerative disease of the central nervous system\u201d (2013, p.29)\u2013anthropological research regarding the the cultural mortuary rites within the Okapa region of Papua New Guinea have linked the newfound disease \u2018Kuru\u2019 to post-mortem consumption of human remains (Radford &amp; Scragg, 2013; Collinge Et al., 2006).<\/p>\n<p>Local oral histories collected during the first investigations by these researchers in the 1950s traced the earliest cases back to the 1920s, with detailed case histories. Epidemiological data revealed a strong correlation between the spread of Kuru and participation in mortuary feasts, in which the deceased were ritually consumed as part of funerary rites (2006, p. 2070). From 1957 to 2004, over 2,700 cases were reported, with mortality peaking at over 200 deaths annually in the late 1950s (2006, p.2070); however, following the cessation of cannibalism in the easly 1960s due to governmental efforts, natural transmission of the disease has stopped, dropping the death toll dramatically, with the \u201clast three single cases reported in 2005, 2007, and 2009\u201d (Radford &amp; Scragg, p.48).<\/p>\n<\/div>\n<h3 class=\"import-Normal\"><strong>The Lasting Gift of Neanderthals: Tantalizing New Directions for Resear<\/strong><strong>ch<\/strong><\/h3>\n<p class=\"import-Normal\">Examining the more recent time period in which Neanderthals lived and the extensive excavations completed across Europe allows for a much more complete archaeological record from this time period. Additionally, the increased cultural complexity such as complex tools and ritual behaviours expressed by Neanderthals left a more detailed record than previous hominins. Intentional burials enhanced preservation of the dead and potentially associated ritual behaviours. Such evidence allows for a more complete and nuanced picture of this species.<\/p>\n<figure style=\"width: 424px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image15-5.png\" alt=\"Museum exhibition of life-sized Neanderthal figure.\" width=\"424\" height=\"469\" \/><figcaption class=\"wp-caption-text\">Figure 12.12: Artistic reconstruction of Neanderthal at The Natural History Museum in Vienna, Austria. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Homo_neanderthalensis,_The_Natural_History_Museum_Vienna,_20210730_1223_1272.jpg\">Homo neanderthalensis, The Natural History Museum Vienna, 20210730 1223 1272<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Jakubhal\">Jakub Ha\u0142un<\/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\">Additional analyses are possible on many Neanderthal finds, due to increased preservation of bone, the amount of specimens that have been uncovered, and the recency in which Neanderthals lived. We should be cautious, however, to consider the potential bias of many Neanderthal sites. Overwhelmingly, Neanderthal skeletons are found complete, with injuries or evidence of disease in caves. Does this mean all Neanderthals lived a tough, disease-wrought life? Probably not. It does, however, indicate that the sick were cared for by others, and that they lived in environments that preserved their bodies incredibly well. These additional studies include the examination of dental calculus and even DNA analysis. While limited, samples of Neanderthal DNA have been successfully extracted and analysed. Research thus far has identified specific genetic markers that show some Neanderthals were light-skinned and probably red-haired with light eyes. Genetic analyses, different from the typical hominin reconstruction done with earlier species, allow scientists to further investigate soft tissue markers of Neanderthals and other more recent hominin species. These studies offer striking conclusions regarding Neanderthal traits, their physical appearance, and their culture, as reflected in these artists\u2019 reconstructions (Figure 12.12).<\/p>\n<figure style=\"width: 258px\" class=\"wp-caption alignright\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image17-4.png\" alt=\"Photograph of Dr. Svante P\u00e4\u00e4bo in a blue suit and red tie.\" width=\"258\" height=\"368\" \/><figcaption class=\"wp-caption-text\">Figure 12.13: Nobel Prize winner (2022) and pioneer in paleogenomic research, Dr. Svante P\u00e4\u00e4bo. Credit: <a href=\"https:\/\/upload.wikimedia.org\/wikipedia\/commons\/2\/26\/Professor_Svante_Paabo_ForMemRS_%28cropped%29.jpg\">Professor Svante Paabo ForMemRS (cropped)<\/a> by Duncan.Hull 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\">Dr. Svante P\u00e4\u00e4bo (Figure 12.13), of the Max Planck Institute for Evolutionary Anthropology, has been at the forefront of much of this new research, largely in the form of genomic studies (The Nobel Prize 2022). Awarded the Nobel Prize for Physiology or Medicine in 2022, P\u00e4\u00e4bo is known primarily for his work with ancient DNA. He has successfully sequenced mitochondrial DNA (mtDNA) as well as the entire Neanderthal genome from nuclear DNA. His genomic work has led to the realization that Denisovans are genetically distinct from Neanderthals, as well as the recent identification of a Neanderthal father and teenage daughter, which he discovered by looking for unique DNA markers in the fossil record. Additionally, P\u00e4\u00e4bo\u2019s genomic work has provided researchers with additional lines of evidence regarding the connections between hominin fossils (such as Neanderthals) and modern people, their time of divergence, and current genetic overlap. The work of P\u00e4\u00e4bo has even formalized a new field of study within anthropology\u2014paleogenomics. To stay up to date with Dr. Svante P\u00e4\u00e4bo\u2019s work, be sure to follow his <a href=\"https:\/\/www.eva.mpg.de\/genetics\/neandertals-and-more\/overview\/\">lab\u2019s website<\/a>.<\/p>\n<h3 class=\"import-Normal\"><strong>Neanderthal Culture: Communicating through Speech<\/strong><\/h3>\n<p class=\"import-Normal\">To successfully live in groups and to foster cultural innovations, Neanderthals would have required at least a basic form of communication in order to function, possibly using a speech-based communication system. The challenge with this line of research is that speech, of course, is not preserved, so indirect evidence must be used to support this conclusion. It is thought that Neanderthals would have possessed some basic speech, as evidenced from a variety of sources, including throat anatomy and genetic evidence (Lieberman 1971). There is only one bone in the human body that could demonstrate if a hominin was able to speak, or produce clear vocalizations like modern humans, and that is the hyoid, a U-shaped bone that is found in the throat and is associated with the ability to precisely control the vocal cords. Very few hyoid bones have been found in the archaeological record; however, a few have been uncovered in Neanderthal burials. The shape of the Neanderthal hyoid is nearly identical to that of modern humans, pointing to the likelihood that they had the same vocal capabilities as modern humans. In addition, geneticists have uncovered a mutation present in both modern humans and Neanderthals\u2014the FOXP2 gene\u2014that is possibly linked to the ability to speak. However, other scientists argue that we cannot make sweeping conclusions that the FOXP2 gene accounts for speech due to small sample size. Finally, scientists have also pointed to the increasingly complex cultural behaviour of Neanderthals as a sign that symbolic communication, likely through speech, would have been the only way to pass down the skills needed to make, for example, a Levallois blade or to position a body for intentional burial.<\/p>\n<h3 class=\"import-Normal\"><strong>Neanderthal Intelligence<\/strong><\/h3>\n<p class=\"import-Normal\">One of the enduring questions about Neanderthals centres on their intelligence, specifically in comparison to modern humans. Brain volume indicates that Neanderthals certainly had a large brain, but it continues to be debated if Neanderthals were of equal intelligence to modern humans. Remember, creatures with larger body sizes tend to have larger brains; however, scaling of the brain is not always associated with greater intelligence (Alex 2018). Brain volume (cranial capacity), cultural complexity, tool use, and compassion toward their kind all point to an increase in intellect among Neanderthals when compared to previous hominins.<\/p>\n<p class=\"import-Normal\">Yet, new research is suggesting additional differences between Neanderthal brains and our own. For example, Euluned Pearce and colleagues (2013), from the University of Oxford, noted the frontal lobes of Neanderthals and modern humans are almost identical. However, Neanderthals had a larger visual cortex\u2014the portion of the brain involved in processing visual information. This would have left Neanderthals with less brain tissue for other functions, including those that would have aided them in dealing with large social groupings, one of the differences that has been suggested to exist between Neanderthals and modern humans. Other differences were found when geneticist John Blangero, from the Texas Biomedical Research Institute, compared data from the Neanderthal genome against data from modern study participants. Blangero and his colleagues (Blangero Et al. 2014) discovered that some Neanderthal brain components were very different, and smaller, than those in the modern sample. Differences were found in areas associated with the processing of information and controlling emotion and motivation, as well as overall brain connectivity. In short, as Blangero stated, \u201cNeanderthals were certainly cognitively adept,\u201d although their specific abilities may have differed from modern humans\u2019 in key areas (qtd. in Wong 2015). This point has been echoed in other recent genetic studies comparing Neanderthal and anatomically modern human brains (el-Showk 2019).<\/p>\n<p class=\"import-Normal\">Finally, scientists are fairly certain that Neanderthal brain development after birth was not the same as that of modern humans. After birth, anatomically modern <em>Homo sapiens <\/em>babies go through a critical period of brain expansion and cognitive development. It appears that Neanderthal babies\u2019 brains and bodies did not follow the same developmental pattern (Smith Et al. 2010; Zollikofer and Ponce de Le\u00f3n 2013). Modern humans enjoy an extended period of childhood, which allows children to engage in imaginative play and develop creativity that fosters cognitive skills. Neanderthals had a more limited childhood, with less development of the creative mind that may have affected their species\u2019 success (Nowell 2016).<\/p>\n<p class=\"import-Normal\">The exact nature of Neanderthal intelligence remains under investigation, however. Some studies disagree with the idea that Neanderthal intelligence had limitations compared to our own, noting the extensive evidence of Neanderthals having limb asymmetry. Their tools also have wear marks indicating that they were hand-dominant. This is further supported by marks on Neanderthal teeth that demonstrate hand dominance. The Neanderthal \u201cstuff-and-cut method\u201d of eating, noted by David Frayer and colleagues (Frayer Et al. 2012), would have seen Neanderthals hold a piece of meat in their teeth, while pulling it taut with one hand, and then using the other hand, their dominant one, to cut the meat off of the larger slab being held in their teeth. When looking at 17 Neanderthals and their tooth wear, only two do not show markings associated with a right-hand dominant individual eating in this manner. Further, it has been established that favouring the right hand is a key marker between modern humans and chimpanzees, and that handedness also relates to language development, in the form of bilateral brain development. That Neanderthals likely were hand-dominant suggests they had an indicator of bilateral brain development and a precondition for human speech.<\/p>\n<h2 class=\"import-Normal\">The Middle Stone Age: Neanderthal Contemporaries in Africa<\/h2>\n<p class=\"import-Normal\">While Neanderthals made their home on and adapted to the European and Asian continents, evidence of fossil humans in Africa show they were also adapting to their local environments. These populations in Africa exhibit many more similarities to modern humans than Neanderthals, as well as overall evolutionary success. While the African fossil sample size is smaller and more fragmentary than the number of Neanderthal specimens across Europe and Asia, the African sample is interesting in that it represents a longer time period and larger geographical area. This group of fossils\u2014often represented by the name \u201cMiddle Stone Age,\u201d or MSA\u2014dates to between 300,000 and 30,000 years ago across the entire continent of Africa. As with Archaic <em>Homo sapiens<\/em>, there is much variability seen in this African set of fossils. There are also a few key consistent elements: none of them exhibit Neanderthal skeletal features; instead, they demonstrate features that are increasingly consistent with anatomically modern <em>Homo sapiens<\/em>.<\/p>\n<p class=\"import-Normal\">Similarities to Neanderthals and MSA contemporaries in Africa are seen, however, in their behavioural adaptations, including stone tools and other cultural elements. The tools associated with the specimens living in Africa during this time period are, like their physical features, varied. In some parts of Africa, namely Northern Africa, stone tools from this time so closely resemble Neanderthal tools that they are classified as Mousterian. In sub-Saharan Africa, the stone tools associated with these specimens are labeled as MSA. Some scholars argue that these could also be a type of Mousterian tools, but they are still typically subdivided based on geographical location.<\/p>\n<p class=\"import-Normal\">Recall that Mousterian tools were much more advanced than their Acheulean predecessors in terms of how the stone tools were manufactured, the quality of the stones used, and the ultimate use of the tools that were made. In addition, recent evidence suggests that MSA tools may also have been heat treated\u2014to improve the quality of the stone tool produced (Stolarczyk and Schmidt 2018). Evidence for heat treating is seen not only through advanced analysis of the tool itself but also through the residue of fires from this time period. Fire residues show a shift over time from small, short fires fuelled by grasses (probably intended for cooking) to larger, more intensive fires that required the exploitation of dry wood, exactly the type of fire that would have been needed for heat treating stone tools (Esteban Et al. 2018).<\/p>\n<p class=\"import-Normal\">Other cultural elements seen with MSA specimens include the use of marine (sea-based) resources for their diet (Parkington 2003), manufacture of bone tools, use of adhesive and compound tools (e.g., hafted tools), shell bead production, engraving, use of pigments (such as ochre), and other more advanced tool-making technology (e.g., microlithics). While many of these cultural elements are also seen to a limited extent among Neanderthals, developments at MSA sites appear more complex. This MSA cultural expansion may have been a response to climate change or an increased use of language, complex communication, and\/or symbolic thought. Others have suggested that the MSA cultural expansion was due to the increase of marine resources in their diet, which included more fatty acids that may have aided their cognitive development. Still others have suggested that the increased cultural complexity was due to increased interaction among groups, which spurred competition to innovate. Recent studies suggest that perhaps the best explanation for the marked cultural complexity of MSA cultural artifacts is best explained by the simple fact that they lived in diverse habitats (Kandel Et al. 2015). This would have necessitated a unique set of cultural adaptations for each habitat type (for example, specialized marine tools would have been needed along coastal sites but not at inland locations). Simply put, the most useful adaptation of MSA was their flexibility of behaviour and adaptability to their local environment. As noted previously in this chapter, flexibility of behaviour and physical traits, rather than specialization, seems to be a feature that was favoured in hominin evolution at this time.<\/p>\n<h2 class=\"import-Normal\">Where Did They Go? The End of Neanderthals<\/h2>\n<p class=\"import-Normal\">While MSA specimens were increasingly successful and ultimately transitioned into modern <em>Homo sapiens<\/em>, Neanderthals disappeared from the fossil record by around 40,000 years ago. What happened to them? We know, based on genetics, that modern humans come largely from the modern people who occupied Africa around 300,000 to 100,000 years ago, at the same time that Neanderthals were living in northern Europe and Asia. As you will learn in Chapter 13, modern humans expanded out of Africa around 150,000 years ago, rapidly entering areas of Europe and Asia inhabited by Neanderthals and other Archaic hominins. Despite intense interest and speculation in fictional works about possible interactions between these two groups, there is very little direct evidence of either peaceful coexistence or aggressive encounters. It is clear, though, that these two closely related hominins shared Europe for thousands of years, and recent DNA evidence suggests that they occasionally interbred (Fu Et al. 2015). Geneticists have found traces of Neanderthal DNA (as much as 1% to 4%) in modern humans of European and Asian descent not present in modern humans from sub-Saharan Africa. This is indicative of limited regional interbreeding with Neanderthals.<\/p>\n<p class=\"import-Normal\">While some interbreeding likely occurred, as a whole, Neanderthals did not survive. What is the cause for their extinction? This question has fascinated many researchers and several theories have been suggested, including:<\/p>\n<ul>\n<li class=\"import-Normal\">At the time that Neanderthals were disappearing from the fossil record, the climate went through both cooling and warming periods\u2014each of which posed challenges for Neanderthal survival (Defleur and Desclaux 2019; Staubwasser Et al. 2018). It has been argued that as temperatures warmed, large-bodied animals, well adapted to cold weather, moved farther north to find colder environments or faced extinction. A shifting resource base could have been problematic for continued Neanderthal existence, especially as additional humans, in the form of modern <em>Homo sapiens<\/em>, began to appear in Europe and compete for a smaller pool of available resources.<\/li>\n<li class=\"import-Normal\">It has been suggested that the eruption of a European volcano 40,000 years ago could have put a strain on available plant resources (Golovanova Et al. 2010). The eruption would have greatly affected local microclimates, reducing the overall temperature enough to alter the growing season.<\/li>\n<li class=\"import-Normal\">Possible differences in cognitive development may have limited Neanderthals in terms of their creative problem solving. As much as they were biologically specialized for their environment, the nature of their intelligence might not have offered them the creative problem-solving skills to innovate ways to adapt their culture when faced with a changing environment (Pearce Et al. 2013).<\/li>\n<li class=\"import-Normal\">CRISPR gene-editing technology has been used in studies to evaluate potential differences between human and Neanderthal brains, based on differences in the genetic code. Potential differences include a Neanderthal propensity for mutations related to brain development that could account for more rapid brain development, maturation, synapse misfires, and less-orderly neural processes (Mora-Berm\u00fadez Et al. 2022; Trujillo Et al. 2021). Fundamental differences in brain function at the cellular level may account for the differential survival rates of Neanderthal and modern human populations.<\/li>\n<li class=\"import-Normal\">There is evidence that suggests reproduction may have posed challenges for Neanderthals. Childbirth was thought to have been at least as difficult for female Neanderthals as anatomically modern <em>Homo sapiens<\/em> (Weaver and Hublin 2009). Female Neanderthals may have become sexually mature at an older age, even older than modern humans. This delayed maturation could have kept the Neanderthal population size small. A recent study has further suggested that male Neanderthals might have had a genetic marker on the Y chromosome that could have caused incompatibility between the fetus and mother during gestation; this would have had severe consequences for birth rate and survival (Mendez Et al. 2016). Even a small but continuous decrease in fertility would have been enough to result in the extinction of Neanderthals (Degioanni Et al. 2019).<\/li>\n<li class=\"import-Normal\">As mentioned above, the end of Neanderthal existence overlaps with modern human expansion into northern Europe and Asia. There is no conclusive direct evidence to indicate that Neanderthals and modern humans lived peacefully side by side, nor that they engaged in warfare, but by studying modern societies and the tendencies of modern humans, it has been suggested that modern humans may not have warmly embraced their close but slightly odd-looking cousins when they first encountered them (Churchill Et al. 2009). Nevertheless, direct competition with modern humans for the same resources may have contributed to the Neanderthals\u2019 decline (Gilpin Et al. 2016); it may also have exposed them to new diseases, brought by modern humans (Houldcroft and Underdown 2016), which further decimated their population. Estimates of energy expenditures suggest Neanderthals had slightly higher caloric needs than modern humans (Venner 2018). When competing for similar resources, the slightly greater efficiency of modern humans might have helped them experience greater success in the face of competition\u2014at a cost to Neanderthals.<\/li>\n<\/ul>\n<p class=\"import-Normal\">It has been suggested that the Neanderthal populations were fairly small to begin with (estimated between 5,000 and 70,000 individuals; Bocquet-Appel and Degioanni 2013), one or a combination of these factors could have easily led to their demise. As more research is conducted, we will likely get a better picture of exactly what led to Neanderthal extinction.<\/p>\n<h2 class=\"import-Normal\">Denisovans<\/h2>\n<figure style=\"width: 353px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image16-4.png\" alt=\"Small fossilized finger bone sitting atop a chalk outline of hand bones.\" width=\"353\" height=\"235\" \/><figcaption class=\"wp-caption-text\">Figure 12.14: Reproduction of Denisovan finger bone. Credit: <a href=\"https:\/\/upload.wikimedia.org\/wikipedia\/commons\/6\/6f\/Denisova_Phalanx_distalis.jpg\">Denisova Phalanx distalis<\/a> (image from Museum of Natural Sciences, Brussels, Belgium) by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Thilo_Parg\">Thilo Parg<\/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\">While Neanderthals represent one regionally adapted branch of the Archaic <em>Homo sapiens <\/em>family tree, recent discoveries in Siberia and the Tibetan Plateau surprised paleoanthropologists by revealing yet another population that was contemporary with Archaic <em>Homo sapiens<\/em>, Neanderthals, and modern <em>Homo sapiens<\/em>. The genetic analysis of a child\u2019s finger bone (Figure 12.14) and an adult upper third molar (Figure 12.15) from Denisova Cave in the Altai Mountains in Siberia by a team including Svante P\u00e4\u00e4bo discovered that the mitochondrial and nuclear DNA sequences reflected distinct genetic differences from all known Archaic populations. Dubbed \u201cDenisovans\u201d after the cave in which the bones were found, this population is more closely related to Neanderthals than modern humans, suggesting the two groups shared an ancestor who split from modern humans first, then the Neanderthal-Denisovan line diverged more recently (Reich Et al. 2010).<\/p>\n<figure style=\"width: 227px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image6-4-1.png\" alt=\"Molar tooth with wear, large surface area, and large roots.\" width=\"227\" height=\"341\" \/><figcaption class=\"wp-caption-text\">Figure 12.15: Reproduction of Denisovan molar. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Denisova_Molar.jpg\">Denisova Molar<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Thilo_Parg\">Thilo Parg<\/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\">Denisovans share up to 5% of their DNA with modern Melanesians, aboriginal Australians, and Polynesians, and 0.2% of their DNA with other modern Asian populations and Native Americans. Additional studies have suggested one (Vernot Et al. 2018) or two (Browning Et al. 2018) separate points of time when interbreeding occurred between modern humans and Denisovans.<\/p>\n<p class=\"import-Normal\">Genetic analysis reveals that Denisovans (potentially three distinct populations) had adaptations for life at high altitudes that prevented them from developing altitude sickness and hypoxia in extreme environments such as Tibet, where the average annual temperature is close to 0\u2103 and the altitude is more than a kilometre (about 4,000 feet) above sea level. Through protein analysis of a jawbone, one study (Chen Et al. 2019) has placed Denisovans in Tibet as early as 160,000 years ago. Genetic evidence of interbreeding has linked modern Tibetan populations with Denisovans 30,000 to 40,000 years ago, which implies that the unique high-altitude adaptations seen in modern Tibetans may have originated with Denisovans (Huerta-Sanchez Et al. 2014).<\/p>\n<p class=\"import-Normal\">Other research suggests tantalizing new directions regarding Denisovans. Stone tools similar to those found in Siberia have been uncovered in the Tibetan plateau suggesting a connection between the Denisovan populations in those two areas (Zhang Et al. 2018). The molar of a young girl, possibly Denisovan, has been found in Laos and shows strong similarities to specimens from China (Demeter Et al. 2022). And DNA sequencing from discoveries in the Denisova Cave have yielded a genome that has been interpreted as the first-generation offspring of a Denisovan father and Neanderthal mother (Slon Et al. 2018). While this research is not yet conclusive and is still being interpreted, exciting new possibilities are being revealed. To stay up-to-date with new discoveries, consider following organizations such as the <a href=\"https:\/\/www.facebook.com\/smithsonian.humanorigins\/\">Smithsonian\u2019s Human Origins Program<\/a> on social media.<\/p>\n<div class=\"textbox\" style=\"background: var(--lightblue)\">\n<h2>Pleistocene Hominins in East Asia: Homo longi and Yunxian<\/h2>\n<p>New fossil discoveries, along with advanced molecular analysis, are prompting re-evaluations of the complex evolutionary dynamics of non-sapiens Homo in the middle-late Pleistocene.<\/p>\n<h3>Homo longi<\/h3>\n<p>The Harbin cranium, often referred to as Homo longi, represents one of the most remarkable Middle Pleistocene human fossils discovered in East Asia. It is currently carbon-dated at 146 ka, allowing recent scholarship to use the Harbin cranium to argue for a distinct clade of hominins that evolved independently in Eastern Asia, separate from Neanderthals and modern humans (Ni et al., 2021). This clade is argued to include Dali, Jinniushan, Hualongdong, Xiahe, and Harbin, based on phylogenetic analyses using parsimony criteria and Bayesian tip-dating, forming a third East Asian lineage and a sister group to the Homo sapiens lineage.<\/p>\n<p>The Harbin cranium exhibits a unique combination of traits, likely representing a species of Homo distinct from other Middle-to-Late Pleistocene taxa, including H. sapiens, H. Neanderthalensis, and H. Heidelbergensis\/H. Rhodesiensis. Its overall size is exceptional, setting it apart from nearly all other known fossils; however, the proportions of the cranial vault align closely with those of other large late archaic Homo species, with an estimated brain volume of approximately 1,420 mL. Despite its considerable breadth, the face is relatively low in height and exhibits a zygomaxillary shape similar to that seen in H. sapiens and H. antecessor, a feature also observed in other Middle Pleistocene Chinese fossils from Dali and Jinniushan. Additionally, the face is attached to the braincase with reduced prognathism, forming a more perpendicular angle as in modern humans, in contrast to the pronounced midfacial prognathism of Neanderthals.<\/p>\n<h3>Yunxian, China: Possible cladistic origins<\/h3>\n<p>In 2025, researchers released findings that expanded the diversity and relationships of Homo lineages in the Pleistocene, based on an analysis of fossils found in 1989\/90. Two skull fossils were found near the Hanjiang River in Hubei province, China, and were named the Yunxian 1 and Yunxian 2 crania\u2014A third cranium has recently been discovered in the same sediment layer as the two previous finds (Tu et al., 2026). The first 2 specimens were in poor condition, fragmented and distorted; however, using recently developed techniques such as high-resolution computed tomography (CT) scans, image segmentation, and digital repositioning of fragments, the researchers incorporated elements from the Yunxian 1 crania with the less damaged Yunxian 2 cranium. The Yunxian 2 cranium is a mosaic of primitive and derived traits. It retains ancient characteristics seen in earlier hominins, a thick supraorbital torus, a broad palate, a long and low vault in lateral view, and a receding frontal contour; its frontal lobe remains low and narrow, similar to H. erectus fossils. Conversely, it lacks the strongly angulated occipital bone found in H. erectus as well as the protruding occipital region or central suprainiac fossa typical of Neanderthals. The big surprise is that the zygomaxillary region is transversely flat and faces anteriorly, a feature comparable to the Hualongdong and Harbin fossils and even H. sapiens. This intermediate morphology places Yunxian 2 in a distinct position, linking archaics like H. erectus with later Asian hominins such as Harbin, Dali, and Jinniushan. Phylogenetic and morphometric analyses confirm that Yunxian 2 represents an early member of the Asian H.longi Clade. The reconstructed Yunxian 2 cranium represents a major achievement in paleoanthropology, digitally restoring a badly damaged fossil from beyond the Middle Pleistocene. In fact, recent discussion suggests an even earlier date for the Yunxian fossils at approximately 1.77 Ma (Tu et al., 2026). In their analysis, they identify the Yunxian fossils as H.erectus, and the site as Early Pleistocene. Over 500 artifacts have been recovered from the site (cores, choppers, and flakes) typical of the Early Paleolithic. Nevertheless, the mosaic morphology of the Yunxian skulls reveals characteristics within the range of both H. erectus and the Harbin skull (among others), which may position these finds as being very close to the origins and diversification of the longi clade, as well as preserving transitional features close to the sapiens clade\u00a0 (Feng et al., 2025, p.1323).<\/p>\n<p>The expanding fossil and genomic record of Archaic Homo has profoundly reshaped our understanding of Pleistocene human evolution. Discoveries, from an increasing variety of fossils found in eastern Asia, to advances in genomic research, demonstrate that multiple contemporaneous lineages once coexisted, interacted, diverged and, at times, interbred. These findings challenge earlier linear models of human ancestry and instead point to a dynamic evolutionary landscape marked by regional diversification, intermittent contact, and complex patterns of gene flow.<\/p>\n<\/div>\n<p>&nbsp;<\/p>\n<h2 class=\"import-Normal\">How Do These Fit In? <em>Homo naledi<\/em> and <em>Homo floresiensis<br \/>\n<\/em><\/h2>\n<p class=\"import-Normal\">Recently, some fossils have been unearthed that have challenged our understanding of the hominin lineage. The fossils of <em>Homo <\/em><em>naledi<\/em> and <em>Homo <\/em><em>floresiensis<\/em> are significant for several reasons but are mostly known for how they don\u2019t fit the previously held patterns of hominin evolution. While we examine information about these species, we ask you to consider the evidence presented in this chapter and others to draw your own conclusions regarding the significance and placement of these two unusual fossil species in the hominin lineage.<\/p>\n<h3 class=\"import-Normal\"><strong>Homo <\/strong><strong>naledi<\/strong><\/h3>\n<figure style=\"width: 347px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image12-5.png\" alt=\"A nearly complete skeleton surrounded by off-white bone fragments on a black table.\" width=\"347\" height=\"390\" \/><figcaption class=\"wp-caption-text\">Figure 12.16: A sample of some of the 1,550 bones found representing Homo naledi. Credit: <a href=\"https:\/\/elifesciences.org\/articles\/09560#fig1\">Dinaledi skeletal specimens (Figure 1)<\/a> by <a href=\"https:\/\/elifesciences.org\/articles\/09560\">Berger et al. 2015<\/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\">In 2013 recreational spelunkers uncovered a collection of bones deep in a cave network in Johannesburg, South Africa. The cave system, known as Rising Star, had been well documented by other cavers; however, it appears few people had ever gone as far into the cave as these spelunkers did. Lee Berger, paleoanthropologist at University of Witwatersrand, in Johannesburg, immediately put out a call for what he termed \u201cunderground astronauts\u201d to begin recovery and excavation of the fossil materials. Unlike other excavations, Berger and most other paleoanthropologists would not be able to access the elusive site, as it was incredibly difficult to reach, and at some points there was only eight inches of space through which to navigate. The underground astronauts, all petite women anthropologists, were the only ones who were able to access this remarkable site. Armed with small excavation tools and a video camera, which streamed the footage up to the rest of the team at the surface, the team worked together and uncovered a total of 1,550 bones, representing at least 15 individuals, as seen in Figure 12.16. Later, an additional 131 bones, including an almost-complete cranium, were found in a nearby chamber of the cave, representing three more individuals (Figure 12.17). Berger called in a team of specialists to participate in what was dubbed \u201cPaleoanthropology Summer Camp.\u201d Each researcher specialized in a different portion of the hominin skeleton. With various specialists working simultaneously, more rapid analysis was possible of <em>Homo <\/em><em>naledi<\/em> than most fossil discoveries.<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 534px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image2-4.png\" alt=\"Photograph of four different views of the LES1 Homo naledi skull set against a black background.\" width=\"534\" height=\"599\" \/><figcaption class=\"wp-caption-text\">Figure 12.17: Several angles of the nearly complete LES1 Homo naledi skull. Credit: <a href=\"https:\/\/elifesciences.org\/articles\/24232#fig5\">LES1 Cranium (Figure 5)<\/a> by <a href=\"https:\/\/elifesciences.org\/articles\/24232\">Hawks et al. 2017<\/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\">While access to the site, approximately 80 m from any known cave entrance or opening, was treacherous for researchers, it must have been difficult for <em>Homo <\/em><em>naledi<\/em> as well. The route included moving through a portion that is just 25 cm wide at some points, known as \u201cSuperman\u2019s Crawl.\u201d The only way to get through this section is by crawling on your stomach with one arm by your side and the other raised above your head. Past Superman\u2019s Crawl, a jagged wall known as the Dragon\u2019s Back would have been very difficult to traverse. Below that, a narrow vertical chute would have eventually led down to the area where the fossils were discovered. While geology changes over time and the cave system likely has undergone its fair share, it is not likely that these features arose after <em>Homo <\/em><em>naledi<\/em> lived (Dirks Et al. 2017). This has made scientists curious as to how the bones ended up in the bottom of the cave system in the first place. It has been suggested that <em>Homo <\/em><em>naledi<\/em> deposited the bones there, one way or another. If <em>Homo <\/em><em>naledi<\/em> did deposit the bones, either through random disposal or intentional burial, this raises questions regarding their symbolic behaviour and other cultural traits, including the use of fire, to access a very dark cave system. Another competing idea is that a few individuals may have entered the cave system to escape a predator and then got stuck. To account for the sheer number of fossils, this would have had to happen multiple times.<\/p>\n<p class=\"import-Normal\">The features of <em>Homo <\/em><em>naledi<\/em> are well-documented due to the fairly large sample, which represents individuals of all sexes and a wide range of ages. The skull shape and features are very much like other members of the genus <em>Homo<\/em>\u2014including a sagittal keel and large brow, like <em>Homo <\/em><em>erectus<\/em>, and a well-developed frontal lobe, similar to modern humans\u2014yet the brain size is significantly smaller than its counterparts, at approximately 500 cc (560 cc for males and 465 cc for females). The teeth also exhibit features of later members of the genus <em>Homo<\/em>, such as Neanderthals, including a reduction in overall tooth size. <em>Homo <\/em><em>naledi<\/em> also had unique shoulder anatomy and curved fingers, indicating similarities to tree-dwelling primates, which is very different from any other hominin yet found. Perhaps the greatest shock of all is that <em>Homo <\/em><em>naledi<\/em> has been dated to 335,000 to 236,000 years ago, placing it as a contemporary to modern <em>Homo sapiens,<\/em> despite its very primitive features. An additional specimen of a child, found in 2021, not only shares many of the unique features found in the adult specimen but will also add insight into the growth and development of individuals of this species (Brophy Et al. 2021).<\/p>\n<h3 class=\"import-Normal\"><strong>Homo <\/strong><strong>floresiensis<\/strong><\/h3>\n<p class=\"import-Normal\">In a small cave called Liang Bua, on the island of Flores, in Indonesia, a small collection of fossils were discovered beginning in 2003 (Figure 12.18). The fossil fragments represent as many as nine individuals, including a nearly complete female skeleton. The features of the skull are very similar to that of <em>Homo <\/em><em>erectus<\/em>, including the presence of a sagittal keel, an arching brow ridges and nuchal torus, and the lack of a chin (Figure 12.19). <em>Homo <\/em><em>floresiensis<\/em>, as the new species is called, had a brain size that was remarkably small at 400 cc, and recent genetic studies suggest a common ancestor with modern humans that predates <em>Homo <\/em><em>erectus<\/em>.<\/p>\n<figure style=\"width: 606px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image8-3.png\" alt=\"View from inside a large cave with people standing near a dug-out square of dirt.\" width=\"606\" height=\"403\" \/><figcaption class=\"wp-caption-text\">Figure 12.18: Liang Bua Cave on the island of Flores, in Indonesia, where a collection of Homo floresiensis specimens were discovered. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Homo_floresiensis_cave.jpg\">Homo floresiensis cave<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/84301190@N00\">Rosino<\/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: 584px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image9-5.png\" alt=\"Photograph of a gray and off-white cast Homo floresiensis skull.\" width=\"584\" height=\"584\" \/><figcaption class=\"wp-caption-text\">Figure 12.19: Homo floresiensis had a brain that was remarkably small at 400 cc. Recent genetic studies suggest a common ancestor with modern humans that predates Homo erectus. Credit: <a href=\"https:\/\/boneclones.com\/product\/homo-floresiensis-skull-BH-033-2\">Homo floresiensis Skull (Flores Skull LB1)<\/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 complete female skeleton, who was an adult, was approximately a metre tall and would have weighed just under 30 kg, which is significantly shorter and just a few kilograms more than the average, modern, young elementary-aged child. A reconstructed comparison between an anatomically modern human and <em>Homo <\/em><em>floresiensis<\/em> can be seen in Figure 12.20. The small size of the fossil has earned the species the nickname \u201cthe Hobbit.\u201d Many questions have been asked about the stature of this species, as all of the specimens found also show evidence of diminutive stature and small brain size. Some explanations include pathology; however, this seems unlikely as all fossils found thus far demonstrate the same pattern. Another possible explanation lies in a biological phenomena seen in other animal species also found on the island, which date to a similar time period. This phenomenon, called <strong>insular dwarfing<\/strong>, is due to limited food resources on an island, which can create a selective pressure for large-bodied species to be selected for smaller size, as an island would not have been able to support their larger-bodied cousins for a long period of time. This phenomenon is the cause of other unique species known to have lived on the island at the same time, including the miniature stegodon, a dwarf elephant species.<\/p>\n<figure style=\"width: 448px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image18-4.png\" alt=\"Black-and-white drawing of a large nude woman and a much smaller man.\" width=\"448\" height=\"611\" \/><figcaption class=\"wp-caption-text\">Figure 12.20: A reconstructed comparison between an anatomically modern human and Homo floresiensis. As an adult, Homo floresiensis was approximately 1 metre tall and would have weighed under 30 kg. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-16\/\">Anatomically modern human and Homo floresiensis (Figure 11.19)<\/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\">There is ongoing research and debate regarding <em>Homo floresiensis<\/em>\u2019 dates of existence, with some researchers concluding that they lived on Flores until perhaps as recently as 17,000 years ago, although they are more often dated to 100,000 to 60,000 years ago. Stone tools from that time period uncovered at the site are similar to other hominin stone tools found on the island of Flores. <em>Homo <\/em><em>floresiensis<\/em> would have hunted a wide range of animals, including the miniature stegodon, giant rats, and other large rodents. Other animals on the island that could have threatened them include the giant komodo dragon. An interesting note about this island chain is that ancestors of <em>Homo <\/em><em>floresiensis<\/em> would have had to traverse the open ocean in order to get there, as the nearest island is almost 10 km away, and there is little evidence to support that a land bridge connecting mainland Asia or Australia to the island would have been present. This separation from the mainland would also have limited the number of other animals, including predators and human species, that would have had access to the island. Anatomically modern <em>Homo sapiens<\/em> arrived on the island around 30,000 years ago and, if some researchers\u2019 later dates for <em>Homo <\/em><em>floresiensis<\/em> are correct, both species may have lived on Flores at the same time. The modern population living on the island of Flores today believes that their ancestors came from the Liang Bua cave; however, recent genetic studies have determined they are not related to <em>Homo <\/em><em>floresiensis<\/em> (Tucci Et al. 2018).<\/p>\n<\/div>\n<h2>Cultural Connections and Contemporary Implications<\/h2>\n<p>The Neanderthals provide an important example of how biological and cultural adaptations cannot always be understood separately. Their large brains and robust, cold-adapted bodies came with significant energetic demands, creating biological challenges that could be addressed through cultural practices. Hunting, tool use, food processing, fire, clothing, and shelter were not simply behaviours that existed alongside their biology; they allowed Neanderthals to respond to the environment and the physiological demands placed on their bodies. In this sense, culture became another means through which Neanderthals adapted to their environment.<\/p>\n<p>Their social behaviours further complicate the distinction between biological survival and cultural adaptation. Evidence of care for injured or vulnerable individuals and intentional burial suggests that their social lives extended beyond immediate individual survival. These social interactions demonstrate that adaptation was not simply a matter of being biologically suited to an environment, but also involved relationships, shared practices, and forms of social care. Although the meaning of these behaviours cannot be fully reconstructed, they challenge the simplistic image of archaic humans as primitive or solely driven by biological instincts.<\/p>\n<p>Ultimately, Neanderthals demonstrate that human adaptation is neither exclusively biological nor exclusively cultural. Their bodies were shaped by the environments in which they lived, while their cultural practices allowed them to navigate and modify those environments. Looking at Neanderthals through a biocultural perspective therefore moves us beyond the idea of the \u201ccaveman\u201d and instead recognizes them as hominins whose survival emerged through the interaction of physiology, environment, technology, and social life.<\/p>\n<h4>Discussion questions<\/h4>\n<ul>\n<li>Why might caring for the sick and injured and practicing burial rites be considered evidence of culture? What do these behaviours reveal about social life?<\/li>\n<li>Can you think of an example of the intertwining of biology and culture in humans today? What does this example reveal about the relationship between biological and cultural adaptation?<\/li>\n<\/ul>\n<p>&nbsp;<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\" style=\"background-color: #ffffff;color: #ffffff\"><span style=\"color: #000000\">Hominin Species Summaries<\/span><\/h2>\n<div style=\"text-align: left\">\n<table class=\"aligncenter\" style=\"width: 468pt\">\n<tbody>\n<tr class=\"Table3-R\" style=\"height: 0\">\n<td class=\"Table3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><strong>Hominin<\/strong><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Archaic <em>Homo sapiens<\/em><\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 0\">\n<td class=\"Table3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><strong>Dates<\/strong><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">600,000\u2013200,000 years ago (although some regional variation)<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 0\">\n<td class=\"Table3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><strong>Region(s)<\/strong><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Africa, Europe, and Asia<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 0\">\n<td class=\"Table3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><strong>Famous discoveries<\/strong><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Broken Hill (Zambia), Atapuerca (Spain)<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 0\">\n<td class=\"Table3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><strong>Brain size<\/strong><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">1,200 cc average<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 0\">\n<td class=\"Table3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><strong>Dentition<\/strong><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Slightly smaller teeth in back of mouth, larger front teeth<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 0\">\n<td class=\"Table3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><strong>Cranial features<\/strong><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Emerging forehead, no chin, projecting occipital region<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 0\">\n<td class=\"Table3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><strong>Postcranial features<\/strong><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Robust skeleton<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 0\">\n<td class=\"Table3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><strong>Culture<\/strong><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Varied regionally, but some continue to use Acheulean handaxe, others adopt Mousterian tool culture<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table3-R\" style=\"height: 0\">\n<td class=\"Table3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\"><strong>Other <\/strong><\/p>\n<\/td>\n<td class=\"Table3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\">Lots of regional variation in this species<\/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: 468pt\">\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\"><strong>Species<\/strong><\/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\"><em>Homo <\/em><em>naledi<\/em><\/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\"><strong>Dates<\/strong><\/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\">335,000\u2013236,000 years ago<\/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\"><strong>Region(s)<\/strong><\/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\">South Africa<\/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\"><strong>Famous discoveries<\/strong><\/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\">Rising Star Cave<\/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\"><strong>Brain size<\/strong><\/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\">500 cc average<\/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\"><strong>Dentition<\/strong><\/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\">Reduced tooth size<\/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\"><strong>Cranial features<\/strong><\/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\">Sagittal keel, large brow, well-developed frontal region<\/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\"><strong>Postcranial features<\/strong><\/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\">Suspensory shoulder<\/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\"><strong>Culture<\/strong><\/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\">unknown<\/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\"><strong>Other<\/strong><\/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\">N\/A<\/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: 468pt\">\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\"><strong>Hominin<\/strong><\/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\">Neanderthals<\/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\"><strong>Dates<\/strong><\/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\">150,000\u201340,000 years ago<\/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\"><strong>Region(s)<\/strong><\/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\">Western Europe, Middle East, and Western Asia only<\/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\"><strong>Famous discoveries<\/strong><\/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\">Shanidar (Iraq), La Chapelle-aux-Saints (France)<\/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\"><strong>Brain size<\/strong><\/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\">1500 cc average<\/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\"><strong>Dentition<\/strong><\/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\">Retromolar gap<\/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\"><strong>Cranial features<\/strong><\/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\">Large brow ridge, midfacial prognathism, large infraorbital foramina, occipital bun<\/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\"><strong>Postcranial features<\/strong><\/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\">Robust skeleton with short and stocky body, increased musculature, barrel chest<\/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\"><strong>Culture<\/strong><\/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\">Mousterian tools often constructed using the Levallois technique<\/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\"><strong>Other<\/strong><\/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\">N\/A<\/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: 468pt\">\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\"><strong>Species<\/strong><\/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\"><em>Homo <\/em><em>floresiensis<\/em><\/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\"><strong>Dates<\/strong><\/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\">100,000\u201360,000 years ago, perhaps as recently as 17,000 years ago<\/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\"><strong>Region(s)<\/strong><\/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\">Liang Bua, island of Flores, Indonesia<\/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\"><strong>Famous discoveries<\/strong><\/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\">\u201cThe Hobbit\u201d<\/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\"><strong>Brain size<\/strong><\/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\">400 cc average<\/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\"><strong>Dentition<\/strong><\/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\">unknown<\/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\"><strong>Cranial features<\/strong><\/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\">Sagittal keel, arching brow ridges, nuchal torus, no chin<\/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\"><strong>Postcranial features<\/strong><\/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\">Very short stature (approximately 3.5 ft.)<\/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\"><strong>Culture<\/strong><\/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\">Tools similar to other tools found on the island of Flores<\/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\"><strong>Other<\/strong><\/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\">N\/A<\/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: 468pt\">\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\"><strong>Hominin<\/strong><\/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\">Denisovans<\/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\"><strong>Dates<\/strong><\/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\">100,000\u201330,000 years ago<\/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\"><strong>Region(s)<\/strong><\/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\">Siberia<\/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\"><strong>Famous discoveries<\/strong><\/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\">Child\u2019s finger bone and adult molar<\/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\"><strong>Brain size<\/strong><\/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\">unknown<\/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\"><strong>Dentition<\/strong><\/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\">Large molars (from limited evidence)<\/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\"><strong>Cranial features<\/strong><\/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\">unknown<\/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\"><strong>Postcranial features<\/strong><\/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\">unknown<\/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\"><strong>Culture<\/strong><\/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\">unknown<\/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\"><strong>Other <\/strong><\/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\">Closely related to Neanderthals (genetically)<\/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\">Summary<\/h2>\n<p class=\"import-Normal\">Research presented in the chapter contributes to why scientists have taken to nicknaming this time period \u201cthe muddle in the middle.\u201d We know that the Middle Pleistocene picks up from <em>Homo erectus <\/em>and ends with the appearance of anatomically modern <em>Homo sapiens<\/em>. While the start and the end are clear, it\u2019s the middle that is messy. As more research is conducted and more data is collected, rather than clarifying our understanding of the hominin lineage during this time period, it only inspires more questions, particularly about the relationships between hominins during this time period, including the oft-misunderstood Neanderthal. Research is painting a more detailed picture of Neanderthal intelligence and both biological and behavioural adaptations. At the same time, their relationship to other Middle Pleistocene hominins, including Denisovans, as well as modern humans, remains unclear.<\/p>\n<p class=\"import-Normal\"><em>Homo<\/em> <em>naledi<\/em> and <em>Homo <\/em><em>floresiensis<\/em> are clear outliers when compared to their contemporary hominin species. Each has surprised paleoanthropologists for both their archaic traits in relatively modern times and their unique combination of traits seen in archaic species and modern humans. While these finds have been exciting, they have also completely upended the assumed trajectory of the human lineage, causing scientists to re-examine assumptions about hominin evolution and what it means to be modern. Add this to the developments being made using ancient DNA, other new fossil discoveries, and other innovations in paleoanthropology, and you see that our understanding of Archaic <em>Homo sapiens<\/em> and others living during this time period is rapidly developing and changing. This is a true testament to the nature of science and the scientific method.<\/p>\n<p class=\"import-Normal\">Clearly, hominins of the Middle Pleistocene are distinct from our species today. Yet, understanding the hominins that directly preceded our species and clarifying the evolutionary relationships between us is important to better understanding our own place in nature.<\/p>\n<h2 class=\"import-Normal\">Review Questions<\/h2>\n<ul>\n<li>What physical and cultural features are unique to Archaic<em> Homo sapiens<\/em>? How are Archaic<em> Homo sapiens<\/em> different in both physical and cultural characteristics from <em>Homo <\/em><em>erectus<\/em>?<\/li>\n<li>Describe the specific changes to the brain and skull first seen in Archaic<em> Homo sapiens<\/em>. Why does the shape of the skull change so dramatically from <em>Homo <\/em><em>erectus<\/em>?<\/li>\n<li>What role did the shifting environment play in the adaptation of Archaic <em>Homo sapiens<\/em>, including Neanderthals? Discuss at least one physical feature and one cultural feature that would have assisted these groups in surviving the changing environment.<\/li>\n<li>What does the regional variation in Archaic <em>Homo sapiens<\/em> represent in terms of the broader story of our species\u2019 evolution?<\/li>\n<li>Describe the issues raised by the discoveries of <em>Homo <\/em><em>naledi<\/em> and <em>Homo <\/em><em>floresiensis<\/em> in the understanding of the story of the evolution of <em>Homo sapiens<\/em>.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<h2 class=\"__UNKNOWN__\">Key Terms<\/h2>\n<div class=\"__UNKNOWN__\">\n<p><strong>Allele<\/strong>: Each of two or more alternative forms of a gene that arise by mutation and are found at the same place on a chromosome.<\/p>\n<p class=\"import-Normal\"><strong>Anthropocentrism<\/strong>: A way of thinking that assumes humans are the most important species and leads to interpreting the world always through a human lens. Species-centric science and thought.<\/p>\n<p class=\"import-Normal\"><strong>Cortex<\/strong>: The outside, or rough outer covering, of a rock. Usually the cortex is removed during the process of stone tool creation.<\/p>\n<p class=\"import-Normal\"><strong>Ethnocentric<\/strong>: Applying negative judgments to other cultures based on comparison to one\u2019s own.<\/p>\n<p class=\"import-Normal\"><strong>Exogenous DNA<\/strong>: DNA that originates from sources outside of the specimen you are trying to sequence.<\/p>\n<p class=\"import-Normal\"><strong>Flexed position<\/strong>: Fetal position, in which the legs are drawn up to the middle of the body and the arms are drawn toward the body centre. Intentional burials are often found in the flexed body position.<\/p>\n<p class=\"import-Normal\"><strong>Foraminifera<\/strong>: Microscopic single-celled organisms with a shell that are common in all marine environments. The fossil record of foraminifera extends back well over 500 million years.<\/p>\n<p class=\"import-Normal\"><strong>Glaciation<\/strong>: A glacial period, or time when a large portion of the world is covered by glaciers and ice sheets.<\/p>\n<p class=\"import-Normal\"><strong>Globular<\/strong>: Round-shaped, like a globe.<\/p>\n<p class=\"import-Normal\"><strong>Grave goods<\/strong>: Items included with a body at burial. Items may signify occupation or hobbies, social status, or level of importance in the community, or they may be items believed necessary for the afterlife.<\/p>\n<p class=\"import-Normal\"><strong>Haft<\/strong>: A handle. Also used as a verb\u2014to attach a handle to an item, such as a stone tool.<\/p>\n<p class=\"import-Normal\"><strong>Infraorbital foramina<\/strong>: Small holes on the maxilla bone of the face that allows nerves and blood to reach the skin.<\/p>\n<p class=\"import-Normal\"><strong>Insular dwarfing<\/strong>: A form of dwarfism that occurs when a limited geographic region, such as an island, causes a large-bodied animal to be selected for a smaller body size.<\/p>\n<p class=\"import-Normal\"><strong>Interglacial<\/strong>: A warmer period between two glacial time periods.<\/p>\n<p class=\"import-Normal\"><strong>Levallois technique<\/strong>: A distinctive technique of stone tool manufacturing used by Archaic <em>Homo sapiens<\/em>, including Neanderthals. The technique involves the preparation of a core and striking edges off in a regular fashion around the core. Then a series of similarly sized pieces can be removed, which can then be turned into different tools.<\/p>\n<p class=\"import-Normal\"><strong>Midfacial prognathism<\/strong>: A forward projection of the nose or the middle facial region. Usually associated with Neanderthals.<\/p>\n<p class=\"import-Normal\"><strong>Mousterian tools<\/strong>: The stone tool industry of Neanderthals and their contemporaries in Africa and Western Asia. Mousterian tools are known for a diverse set of flake tools, which is different from the large bifacial tools of the Acheulean industry.<\/p>\n<p class=\"import-Normal\"><strong>Nasal aperture<\/strong>: The opening for the nose visible on a skull. Often pear- or heart-shaped.<\/p>\n<p class=\"import-Normal\"><strong>Occipital bun<\/strong>: A prominent bulge or projection on the back of the skull, specifically the occipital bone. This is a feature present only on Neanderthal skulls.<\/p>\n<p class=\"import-Normal\"><strong>Ochre<\/strong>: A natural clay pigment mixed with ferric oxide and clay and sand. Ranges in colour from brown to red to orange.<\/p>\n<p class=\"import-Normal\"><strong>Retracted face<\/strong>: A face that is flatter.<\/p>\n<p class=\"import-Normal\"><strong>Retromolar gap<\/strong>: A space behind the last molar and the end of the jaw. This is a feature present only on Neanderthals. It also occurs through cultural modification in modern humans who have had their third molars, or wisdom teeth, removed.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration<\/h2>\n<p><a href=\"https:\/\/www.amnh.org\/exhibitions\/permanent-exhibitions\/anne-and-bernard-spitzer-hall-of-human-origins\">Anne and Bernard Spitzer Hall of Human Origins<\/a>\u2014American Museum of Natural History.<\/p>\n<p>\u201cDawn of Humanity,\u201d PBS documentary, 2015<\/p>\n<p><a href=\"https:\/\/www.ted.com\/talks\/svante_paeaebo_dna_clues_to_our_inner_neanderthal?language=en\">\u201cDNA Clues to Our Inner Neanderthal,\u201d<\/a> TED Talk by Svante P\u00e4\u00e4bo, 2011.<\/p>\n<p>\u201cThe Dirt\u201d Podcast, Episode 30, <a href=\"https:\/\/thedirtpod.com\/episodes\/\/episode-30-the-human-family-tree-shrub-crabgrass-tumbleweed-part-3\">\u201cThe Human Family Tree (Shrub? Crabgrass? Tumbleweed?), Part 3: Very Humany Indeed\u201d<\/a>.<\/p>\n<p class=\"import-Normal\"><a href=\"https:\/\/www.efossils.org\/page\/games-and-activities\">eFossil Games and Activities<\/a><\/p>\n<p>Frank, Rebecca. 2021. \u201cThe Genus Homo.\u201d In <a href=\"https:\/\/explorations.americananthro.org\/index.php\/lab-and-activities-manual\/\"><em>Explorations: Lab and Activity Manual, <\/em><\/a>edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff. Arlington, VA: American Anthropological Association.<\/p>\n<p><a href=\"https:\/\/humanorigins.si.edu\/research\/asian-research-projects\/hobbits-flores-indonesia.\">Hobbits on Flores, Indonesia<\/a> - Smithsonian Human Origins.<\/p>\n<p><a href=\"https:\/\/evolution.berkeley.edu\/evo-news\/lumping-or-splitting-in-the-fossil-record\/\">Lumping or Splitting in the Fossil Record<\/a> - UC Berkeley Understanding Evolution.<\/p>\n<p><a href=\"https:\/\/www.eva.mpg.de\/genetics\/neandertals-and-more\/overview\/\">Neandertals and More<\/a> - Max Planck Institute for Evolutionary Anthropology.<\/p>\n<p><a href=\"https:\/\/www.sapiens.org\/biology\/neanderthal-anatomy\/?fbclid=IwAR2Bcff1GVkTLnbCR58JAWiJzkk-Ell7zL0FUddf1HNAX6RDAZxbqh1zWoI\">Neanderthals: Body of Evidence<\/a> - SAPIENS.<\/p>\n<p>Perash, Rose L., and Kristen A. Broehl. 2021. \u201cHominin Review: Evolutionary Trends.\u201d In <a href=\"https:\/\/explorations.americananthro.org\/index.php\/lab-and-activities-manual\/.\"><em>Explorations: Lab and Activity Manual<\/em><\/a>, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff. Arlington, VA: American Anthropological Association.<\/p>\n<p>Perkl, Bradley. \u201cBrain, Language, Lithics.\u201d In <a href=\"https:\/\/explorations.americananthro.org\/index.php\/lab-and-activities-manual\/.\"><em>Explorations: Lab and Activity Manual<\/em><\/a>, edited by<em>.<\/em> Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff. CC BY-NC. Arlington, VA: American Anthropological Association.<\/p>\n<p><a href=\"https:\/\/humanorigins.si.edu\/evidence\/human-fossils\/shanidar-3-neanderthal-skeleton\">Shanidar 3 - Neanderthal Skeleton<\/a> - Smithsonian Human Origins.<\/p>\n<p><a href=\"https:\/\/humanorigins.si.edu\/evidence\/human-fossils\/species\">Species<\/a> - Smithsonian Human Origins.<\/p>\n<p><a href=\"https:\/\/www.facebook.com\/smithsonian.humanorigins\/\">Smithsonian Human Origins Program Facebook page<\/a> (@smithsonian.humanorigins).<\/p>\n<p><a href=\"https:\/\/www.smithsonianmag.com\/science-nature\/bringing-human-evolution-life-180951155\/\">Paleoartist Brings Human Evolution to Life<\/a> - Elisabeth Dayn\u00e9s.<\/p>\n<h2 class=\"import-Normal\">References<\/h2>\n<p>Ackermann, Rebecca R., Matthew L. Arnold, et al. \u201cHybridization in Human Evolution: Insights from Other Organisms.\u201d Evolutionary Anthropology 28, no. 4 (July 2019): 189\u2013209. <a href=\"https:\/\/doi.org\/10.1002\/evan.21787\">https:\/\/doi.org\/10.1002\/evan.21787<\/a>. <a href=\"https:\/\/pmc.ncbi.nlm.nih.gov\/articles\/PMC6980311\/\">https:\/\/pmc.ncbi.nlm.nih.gov\/articles\/PMC6980311\/<\/a><\/p>\n<p class=\"import-Normal\">Adler, Daniel S., Timothy J. Prindiville, and Nicholas J. Conard. 2003. \u201cPatterns of Spatial Organization and Land Use During the Eemian Interglacial in the Rhineland: New Data from Wallertheim, Germany.\u201d Eurasian Prehistory 1(2): 25\u201378.<\/p>\n<p>Ahlquist, K. D., Mayra M. 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Elliott. 2016. \u201cMultimodal Spatial Mapping and Visualization of Dinaledi Chamber and Rising Star Cave.\u201d <em>South African Journal of Science<\/em> 112 (5\u20136). <a class=\"rId155\" href=\"https:\/\/doi.org\/10.17159\/sajs.2016\/20160032.\">https:\/\/doi.org\/<\/a><a class=\"rId156\" href=\"https:\/\/doi.org\/10.17159\/sajs.2016\/20160032.\">10.17159\/sajs.2016\/20160032.<\/a><\/p>\n<p class=\"import-Normal\">Lieberman, Philip, and Edmund S. Crelin. 1971. \u201cOn the Speech of Neanderthal Man.\u201d <em>Linguistic Inquiry<\/em> 2 (2): 203\u2013222.<\/p>\n<p class=\"import-Normal\">Mendez, Fernando L., G. David Poznik, Sergi Castellano, and Carlos D. Bustamante. 2016. \u201cThe Divergence of Neanderthal and Modern Human Y Chromosomes.\u201d <em>American Journal of Human Genetics<\/em> 98 (4): 728\u2013734.<\/p>\n<p class=\"import-Normal\">Mora-Berm\u00fadez, Felipe, Philipp Kanis, Dominik Macak, Jula Peters, Ronald Naumann, Lei Xing, Mihail Sarov, et al. 2022. \u201cLonger Metaphase and Fewer Chromosome Segregation Errors in Modern Human Than Neanderthal Brain Development.\u201d <em>Science Advances<\/em> 8 (30). <a class=\"rId157\" href=\"https:\/\/doi.org\/10.1126\/sciadv.abn7702.\">https:\/\/doi.org\/10.1126\/sciadv.abn7702.<\/a><\/p>\n<p>Ni, Xijun, Qiang Ji, Wensheng Wu, et al. \u201cMassive Cranium from Harbin in Northeastern China Establishes a New Middle Pleistocene Human Lineage.\u201d The Innovation 2 (June 25, 2021): 100130. <a href=\"https:\/\/doi.org\/10.1016\/j.xinn.2021.100130\">https:\/\/doi.org\/10.1016\/j.xinn.2021.100130<\/a><\/p>\n<p class=\"import-Normal\">Nicholson, Christopher M. 2017. \u201cEemian Paleoclimate Zones and Neanderthal Landscape Use: A GIS Model of Settlement Patterning during the Last Interglacial.\u201d <em>Quaternary International<\/em> 438 (B): 144\u2013157. <a class=\"rId158\" href=\"https:\/\/doi.org\/10.1016\/j.quaint.2017.04.023.\">https:\/\/doi.org\/10.1016\/j.quaint.2017.04.023.<\/a><\/p>\n<p class=\"import-Normal\">Nobel Prize, The. 2022. \u201cPress Release: The Nobel Prize in Physiology or Medicine 2022.\u201d Nobelf\u00f6rsamlingen The Nobel Assembly at Karolinska Institutet. Press release, November 3, 2022. <a class=\"rId159\" href=\"https:\/\/www.nobelprize.org\/prizes\/medicine\/2022\/press-release\/.\">https:\/\/www.nobelprize.org\/prizes\/medicine\/2022\/press-release\/.<\/a><\/p>\n<p class=\"import-Normal\">Nowell, April. 2016. \u201cChildhood, Play and the Evolution of Cultural Capacity in Neanderthals and Modern Humans.\u201d In<em> The Nature of Culture: Based on an Interdisciplinary Symposium \u201cThe Nature of Culture,\u201d T\u00fcbingen, Germany<\/em>, edited by M. N. Haidle, N. J. Conard, and M. Bolus, 87\u201397. Heidelberg, Germany: Springer. <a class=\"rId160\" href=\"blank\">https:\/\/doi.org.10.1007\/978-94-017-7426-0_9<\/a>.<\/p>\n<p class=\"import-Normal\">Parkington, John. 2003. \u201cMiddens and Moderns: Shellfishing and the Middle Stone Age of the Western Cape, South Africa.\u201d <em>South African Journal of Science<\/em> 99 (5\u20136): 243\u2013274.<\/p>\n<p class=\"import-Normal\">Pearce, Eiluned, Chris Stringer, and R. I. M. 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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 src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/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 13.1: 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 analysed 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 12) 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.<\/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 behaviour 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_1683_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 src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image29-2.png\" alt=\"A rounded skull facing a robust skull with sloping forehead.\" width=\"445\" height=\"221\" \/><figcaption class=\"wp-caption-text\">Figure 13.2: 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_1683_1758\"><strong>globular<\/strong>,<\/a> on a modern skull (Lieberman, McBratney, and Krovitz 2002; Neubauer, Hublin, and Gunz 2018; Pearson 2008; Figure 13.2). 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_1683_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 favoured 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 centreline 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 src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 13.3: 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 13.3). 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). A slender frame is believed to be adapted for the efficient long-distance running ability that started with <em>Homo erectus<\/em>. Furthermore, it is argued that 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.<\/p>\n<h3 class=\"import-Normal\"><strong>Behavioural 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 behavioural 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 behavioural 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 13.4). 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 behaviour 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 behaviour. 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 src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 13.4: 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 behaviour. 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 behavioural 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? The key is the set of derived biological traits from the last section. It is theorized that the gracile frame and neurological anatomy allowed modern humans to survive and even flourish in the vastly different environments they encountered. 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_1683_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 13.5 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 class=\"aligncenter\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 class=\"aligncenter\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 class=\"aligncenter\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 13.5a-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><strong>Early Modern <\/strong><strong><em>Homo sapiens\u00a0 <\/em><\/strong> or Early Anatomically Modern Humans<\/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 class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 13.6: 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 13.6). 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 kilometres 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 13.7). 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), favouring categorization with our own species.<\/p>\n<figure style=\"width: 373px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-3.jpg\" alt=\"Replica cranium showing wide brow ridges and gracile face.\" width=\"373\" height=\"373\" \/><figcaption class=\"wp-caption-text\">Figure 13.7: 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_1683_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_1683_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 behavioural modernity was not so developed. No clear signs of art dating back this far have been found either. Other hypotheses not related to behavioural 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 behaviour 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 13.8; 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 class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5-2-1.jpg\" alt=\"Multiple views of shells with holes bored through them.\" width=\"563\" height=\"482\" \/><figcaption class=\"wp-caption-text\">Figure 13.8: 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_1683_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<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_1683_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_1683_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 13.9). 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 behaviour. 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 13.10; Coqueugniot Et al. 2014). Perforated seashells coloured with <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1806\"><strong>ochre<\/strong><\/a>, 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 src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 13.9: 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 src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 13.10 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 colour 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 13.11; 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 src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 13.11: 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 (colour 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 13.12), 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 src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image12-1.jpg\" alt=\"A cave opening amongst a dry wooded region.\" width=\"610\" height=\"458\" \/><figcaption class=\"wp-caption-text\">Figure 13.12: 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<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><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1772\">Sunda<\/a><\/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_1683_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_1683_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_1683_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 src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image18-1-1.jpg\" alt=\"A cranium showing a diagonal sloping forehead.\" width=\"380\" height=\"252\" \/><figcaption class=\"wp-caption-text\">Figure 13.13: 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 colour 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 behaviour, including intentional burial. The bones of Lake Mungo 1 (LM1), an adult female, were crushed repeatedly, coloured 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 13.13).<\/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 behavioural 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<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 in 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. <span style=\"margin: 0px;padding: 0px\">It is suggested that another obstacle was the colder climate, which was incompatible with the biology of modern <em>Homo sapiens<\/em> from Africa, as they were adapted to high temperatures 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_1683_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.<\/p>\n<figure style=\"width: 323px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image11-3.jpg\" alt=\"Robust cranium with a gradually sloping forehead.\" width=\"323\" height=\"323\" \/><figcaption class=\"wp-caption-text\">Figure 13.14: 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 13.14; 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 13.15; 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 src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 13.15: 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, <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1792\"><strong>atlatl<\/strong><\/a> (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 13.16).<\/p>\n<figure style=\"width: 511px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 13.16: 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>.<\/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'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 13.17). 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 class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image24-2-1.jpg\" alt=\"Charcoal painting of a bull seen from the side.\" width=\"605\" height=\"454\" \/><figcaption class=\"wp-caption-text\">Figure 13.17: 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 (colour modified) and is under a <a href=\"https:\/\/whc.unesco.org\/en\/licenses\/6\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<div class=\"textbox shaded\" style=\"background: var(--lightblue)\">\n<h2>Special Topic: Cannibalism and Culture - Mortuary Practices in Modern Homo sapiens<\/h2>\n<p>Within a 2017 publication in the <em>Journal of Archaeological Method and Theory<\/em>, Saladi\u00e9 and Rodr\u00edguez-Hidalgo bring light to traces of early cannibalism in western Eurasia, arguing that context-specific cannibalistic practices were present throughout the Pleistocene and increased notably from the end of the Upper Palaeolithic and onward (Saladi\u00e9 &amp; Rodriguez-Hidalgo, 2017). While early hominins and Neandertals are recognized in this research, the authors highlight the presence of these mortuary practices in a cluster of <em>Homo sapiens<\/em> sites. More recent research uncovers similar findings that back these claims as well, where human bones in Herto Ethiopia, Maszycka Cave Poland, and Gough\u2019s Cave in the United Kingdom show anthropogenic defleshing and other modifications which have been interpreted as cannibalism (Pobiner, Et al. 2023). These findings suggest that cannibalistic behaviours formed a recurring aspect of modern human behaviour in certain ecological and cultural contexts.<\/p>\n<figure id=\"attachment_817\" aria-describedby=\"caption-attachment-817\" style=\"width: 378px\" class=\"wp-caption alignleft\"><img class=\" wp-image-817\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2023\/06\/d_-_briana_pobiner_-_figure_6.jpg\" alt=\"\" width=\"378\" height=\"369\" \/><figcaption id=\"caption-attachment-817\" class=\"wp-caption-text\">Figure 13.18: Close-up photos of three fossil animal specimens from the same area and time horizon as the fossil hominin tibia studied by the research team. These fossils show similar cut marks to those found on the hominin tibia studied. The photos show (a) an antelope mandible, (b) an antelope radius (lower front leg bone) and (c) a large mammal scapula (shoulder blade). Credit: <em data-start=\"617\" data-end=\"635\">23-199D Figure 6<\/em> by Smithsonian\u2019s National Museum of Natural History, from \"<a href=\"https:\/\/www.si.edu\/newsdesk\/releases\/humans-evolutionary-relatives-butchered-one-another-145-million-years-ago\">Humans\u2019 Evolutionary Relatives Butchered One Another 1.45 Million Years Ago<\/a>,\" \u00a9 Smithsonian Institution, used with permission.<\/figcaption><\/figure>\n<p>A significant example comes from the Neolithic levels of Fontbr\u00e9gua Cave in southeastern France, where Paola Villa and colleagues compared clusters of human bones with the remains of wild and domestic animals from the same sediments. The study found that human bodies were butchered, processed, and most likely eaten in a way that parallels animal carcass treatment, including the placement and timing of cut marks, dismemberment sequences, and perimortem fractures to open marrow cavities (Villa, Et al. 1986). As the assemblage comes from a primary depositional context with pristine preservation and careful excavation, the authors suggest that cannibalism is the only satisfactory explanation for the pattern of cut marks and breakage seen on the human bones.<\/p>\n<p>More recent work has furthered this hypothesis for Late Upper Palaeolithic <em>Homo sapiens<\/em>, especially in Magdalenian contexts. In a 2023 study, researchers combined archeological and genetic evidence from fifty-nine Magdalenian sites, concluding that this specific culture shows an unusually high frequency of cannibalistic cases compared to earlier and later hominin groups; so much so that they identify \u201cprimary burial and cannibalism\u201d as the two main mortuary expressions (Marsh &amp; Bello, 2023). Additionally, new analyses from Maszycka Cave in Poland have described cut and broken human bones in patterns consistent with human consumption, reinforcing this cannibalism hypothesis (Marginedas &amp; Saladi\u00e9, 2025). Together, these studies suggest that for some Magdalenian groups, mortuary cannibalism was a habitual way of disposing of the dead, not just a one-off crisis response. At the same time, researchers emphasize that not every modified skeleton indicates blatant consumption of the dead and that ritual or symbolic perspectives must also be considered. Previously noted in chapter 12, Ullrich\u2019s survey of European mortuary practices presents frequent manipulations of corpses from the Palaeolithic through the Hallstatt period. Said manipulations include cut marks, dismemberment, skull fracturing, and marrow extraction (Ullrich, 2005, p. 258), which Ullrich interprets as cannibalistic rites embedded in cult ceremonies rather than everyday subsistence. In the author\u2019s view, some Palaeolithic groups may have believed that consuming members of their community allowed them to take on the strengths,<\/p>\n<figure id=\"attachment_819\" aria-describedby=\"caption-attachment-819\" style=\"width: 265px\" class=\"wp-caption alignright\"><img class=\"wp-image-819\" src=\"http:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/wp-content\/uploads\/sites\/71\/2023\/06\/a_-_briana_pobiner_-_figure_1.jpg\" alt=\"\" width=\"265\" height=\"379\" \/><figcaption id=\"caption-attachment-819\" class=\"wp-caption-text\">Figure 13.19: View of the hominin tibia and magnified area that shows cut marks. Scale = 4 cm. Credit: 23-199A Figure 1 by Jennifer Clark, Smithsonian Institution, from \"<a href=\"https:\/\/www.si.edu\/newsdesk\/releases\/humans-evolutionary-relatives-butchered-one-another-145-million-years-ago\">Humans\u2019 Evolutionary Relatives Butchered One Another 1.45 Million Years Ago<\/a>,\" \u00a9Smithsonian Institution, used with permission.<\/figcaption><\/figure>\n<p>abilities, or even mental aspects of their dead member; the act of cannibalism may have functioned as a form of appropriating physical and mental powers rather than simply obtaining calories. With that, Neolithic assemblages show how careful taphonomic work can distinguish cuts linked to interpersonal violence from those associated with systemic butchery (Marginedas &amp; Saladi\u00e9, 2025). The findings seek to highlight that some <em>Homo sapiens<\/em> populations combined ritual, mortuary, and nutritional motives when processing human remains.<\/p>\n<p>These past and contemporary findings are significant for future anthropology students as they shed light on biological anthropology methodology and interpretation. Archaeologists are able to distinguish between occasions when humans were handled like any other carcass and times when they were handled in more symbolic ways thanks to factors like cut mark orientation, the timing of bone breakage, and direct comparison with animal remains. Readers interested in exploring this topic further should investigate the context-specific motivations for cannibalism, like nutritional stress, warfare, funerary sites, or ritual power appropriation.<\/p>\n<p>Similar archeological techniques have been used to explore behaviours of cannibalism among Indigenous Huron-Wendat populations in 1651 Canada, where human bones exhibit cutmarks, perimortem fractures, and thermal modifications (Spence &amp; Jackson, 2014). These shared taphonomic signatures across continents reveal recurring practices under ecological stress, bridging prehistoric Europe to North American contexts. Engaging with such analytical techniques opens up new ways that archeologists and anthropologists can investigate the variability in human behaviour, from nutritional crises to mortuary rituals.<\/p>\n<\/div>\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_1683_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_1683_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_1683_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 centre: 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 13.20), mastodons, camels, horses, ground sloths, giant tortoises, and\u2014a favourite of researchers\u2014a two-metre-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 class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image3-2-1.jpg\" alt=\"A mammoth model with long curving tusks.\" width=\"242\" height=\"323\" \/><figcaption class=\"wp-caption-text\">Figure 13.20: 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\">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.<\/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. 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 class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image21-4.png\" alt=\"A long stone point with small chips around the edge.\" width=\"254\" height=\"362\" \/><figcaption class=\"wp-caption-text\">Figure 13.21: 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 13.21). 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<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_1683_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 13.22). 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 src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 13.22: 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\">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. 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.<\/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_1683_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_1683_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_1683_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, it has been suggested that 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.<\/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 <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1785\"><strong>bands<\/strong><\/a>, or kin-based groups of around 50 people or less (rarely over 100). A band\u2019s organization would be <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1786\"><strong>e<\/strong><strong>galitarian<\/strong><\/a>, 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 13.23). In varied environments\u2014from savannas to tropical forests, deserts, coasts, and the Arctic circle\u2014people found sustenance needed for survival.<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 565px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image22.jpg\" alt=\"A hunter holding a bow is crouched among dry grass.\" width=\"565\" height=\"377\" \/><figcaption class=\"wp-caption-text\">Figure 13.23: 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 (colour 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\">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 labour increased greatly in poor or unreliable environments. Labour 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\">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.<\/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_1683_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><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1788\">carrying capacity<\/a><\/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><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1789\">Last Glacial Maximum<\/a><\/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 <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1790\"><strong>Younger Dryas<\/strong>.<\/a> 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 labour to farming the limited area. After the Younger Dryas ended, people expanded out of the clusters with their agricultural knowledge (Figure 13.24).<\/p>\n<figure style=\"width: 570px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 13.24: 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. Not only had a growing population may have led to increased competition with other bands, but environments worldwide had shifted to create more uncertainty. As such, it has been proposed that as people in different areas around the world faced this unpredictable situation, they became the independent inventors of agriculture.<\/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, it has been suggested that cultures with no knowledge of each other turned toward intensely farming their local resources (see Figure 13.24).\u00a0 It is proposed that 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). The regions with the most widespread impact in the face of these obstacles became the primary centres of agriculture (Figure 13.25; 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 centre, 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 src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 13.25: 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 (colour 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 neighbouring parts of the world with their domesticates, further expanding the use of this subsistence strategy. From this point, the human species changed from being primarily foragers to primarily agriculturalists with skilled control of their environments. 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><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1793\">sedentarism<\/a><\/strong>. 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.<\/p>\n<figure style=\"width: 359px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image20-1-1.jpg\" alt=\"Multistory buildings surrounding a greek-style plaza.\" width=\"359\" height=\"270\" \/><figcaption class=\"wp-caption-text\">Figure 13.26: 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). It is argued that 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. The surplus of a bountiful harvest also provided insurance for harder times, reducing the risk of famine and bringing change to society as well. With a few farming households producing enough food to feed many others, people could start specializing in roles 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, and over time, the growth of these settlements led to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1794\">urbanization<\/a><\/strong>, the founding of cities that became the foci of human interaction (Figure 13.26).<\/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><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1795\">institutions<\/a><\/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><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1796\">monumental architecture<\/a><\/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 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 centres of mass manufacturing and spurred the rapid development of inventions (Figure 13.27). 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 src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image1-3.jpg\" alt=\"A yellow farm vehicle driving into crops in a field.\" width=\"465\" height=\"310\" \/><figcaption class=\"wp-caption-text\">Figure 13.27: 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 centred 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 favouring 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 labour 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 favour 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><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1798\">aquaculture<\/a><\/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\">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 13.28). 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 src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 13.28: 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 defence 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 colour, this means that populations all around the world could exhibit the whole range of skin colours, 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>Cultural Connections and Contemporary Implications<\/h2>\n<p>The expansion of <em>Homo sapiens<\/em> across the globe and the subsequent and increasingly complex societies they developed have fundamentally altered the relationship between humans and the natural world. Throughout this chapter, we have seen how cultural innovations\u2014agriculture, urbanization, and technological development\u2014enabled humans to modify the environment in ways that no previous hominin could. At this point, the cumulative changes by <em>Homo sapiens<\/em> have irreversibly changed the planet, and some anthropologists and Earth scientists argue that we have entered a new stage of Earth\u2019s history known as the <em>Anthropocene<\/em> (notably the Nobel Laureate Paul J. Crutzen, known as one of the forces in popularizing the term and his part in the <em>Montreal Protocol; <\/em>cf<em>. <\/em>United Nations, 1987; Steffen, 2021<\/p>\n<p>The Anthropocene (or <em>Human Epoch<\/em>) describes a period in which natural processes and human activities have become inseparable, such that the future of Earth's ecosystems and humanity are increasingly dependent on one another (Steffen, 2021). This proposed epoch emphasizes humans\u2019 role as a planetary force capable of reshaping Earth on multiple levels; landscapes are transformed by mining, dam construction, urban expansion, and resource extraction. With this, we are witnessing increasing extinction, the introduction of invasive species, and genetic modification of organisms and unlike earlier periods of human evolution, these changes are occurring simultaneously across the entire planet, affecting ecological systems on a global scale. Although the Anthropocene has been proposed as a new geological epoch beginning in the mid-twentieth century, it has been refused formal adoption within the Geological Time Scale It is generally considered as a part of the Holocene, where agriculture and pastoralism significantly changed the earth despite an argument from a working group in the IPCC (Waters, C 2016).<\/p>\n<p>Nevertheless, the concept of the Anthropocene places modern human societies in positions of unprecedented responsibility. Climate change is not experienced equally, with poverty and social inequality increasing the vulnerability to these challenges, framing the Anthropocene as an environmental concept and a biocultural one as well. Just as agriculture transformed human societies, the choices made by modern populations regarding energy use, technology, and resource management will shape the environments inhabited by future generations and the continued evolution of our species. The Anthropocene therefore reminds us that anthropology and other disciplines aide us in understanding the evolutionary past they are also essential for understanding humanity's role in shaping the future.<\/p>\n<h4>Discussion questions<\/h4>\n<ul>\n<li>How do you see humans shaping the Earth today? Which of these changes do you consider permanent, temporary, harmful, or beneficial?<\/li>\n<li>How might the Anthropocene affect humans as a species? Like other species affected by environmental change, do you think humans could face extinction, or will our ability to modify our environment allow us to adapt?<\/li>\n<li>If humans eventually become a multiplanetary species, would the Anthropocene simply continue beyond Earth? What might this mean for our relationship with the environments we inhabit?<\/li>\n<\/ul>\n<div class=\"textbox\">\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\">Summary<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">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 behavioural 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.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">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.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">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.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">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.<\/p>\n<h2 class=\"import-Normal\">Review Questions<strong><br \/>\n<\/strong><\/h2>\n<ul>\n<li>What are the skeletal and behavioural 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 favourite 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 centres 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 labour-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\">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. 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Cannibalism and burial in the late Upper Palaeolithic: Combining archaeological and genetic evidence. <em>Quaternary Science Reviews<\/em>, 319, 108309. https:\/\/doi.org\/10.1016\/j.quascirev.2023.108309<\/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>Pobiner, B., Pante, M. &amp; Keevil, T. (2023). Early Pleistocene cut marked hominin fossil from Koobi Fora, Kenya. <em>Sci Rep,<\/em> 13, 9896. <a href=\"https:\/\/doi.org\/10.1038\/s41598-023-35702-7\">https:\/\/doi.org\/10.1038\/s41598-023-35702-7<\/a><\/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. 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Cannibalistic rites within mortuary practices from the palaeolithic to middle aged in Europe. Anthropologie (1962-), 43(2\/3), 249\u2013261. <a href=\"http:\/\/www.jstor.org\/stable\/26292739\">http:\/\/www.jstor.org\/stable\/26292739<\/a><\/p>\n<p>United Nations. (n.d.). About Montreal Protocol. UN Environment Programme. <a href=\"https:\/\/www.unep.org\/ozonaction\/who-we-are\/about-montreal-protocol\">https:\/\/www.unep.org\/ozonaction\/who-we-are\/about-montreal-protocol<\/a><\/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>Villa, P., Bouville, C., Courtin, J., Helmer, D., Mahieu, E., Shipman, P., Belluomini, G. &amp; Branca, M. (1986). Cannibalism in the Neolithic. <em>Science<\/em>, 233(4762), 431\u2013437. doi:10.1126\/science.233.4762.431<\/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>Waters, Colin N, et al .The Anthropocene is functionally and stratigraphically distinct from the Holocene <em>Science <\/em>\u00a08 Jan 2016: Vol. 351, Issue 6269.<\/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\">Acknowledgements<\/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>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1683_1712\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1712\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<div class=\"textbox\">\n<p>Learn more about <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/about-the-authors\/\">authors and editors<\/a>,<a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/student-contributors\/\"> Hess' student contributions<\/a>, and <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/versioning-history\/\">versioning history<\/a>.<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\">Learning Objectives<\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li class=\"import-Normal\">Describe the variables that affect primate diets.<\/li>\n<li class=\"import-Normal\">Explain how primates interact with other organisms in their environment.<\/li>\n<li class=\"import-Normal\">Discuss why primates live in groups, types of primate groups, and components of their social systems.<\/li>\n<li class=\"import-Normal\">Describe the reproductive strategies of males and females.<\/li>\n<li class=\"import-Normal\">Explain the ways in which primates communicate.<\/li>\n<li class=\"import-Normal\">Discuss the evidence for primate cultural traditions.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p class=\"import-Normal\">Nonhuman primates (hereafter, 'primates') are a fascinating group of animals, whose similarity to humans can be striking. Because of this similarity, studying primates helps anthropologists to gain insight into how our human ancestors may have behaved. It also allows us to better understand our own behaviour through <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_928\">comparison<\/a><\/strong> (examining similarities and differences) with other primates as well as by comparing different species of primates to one another. In this way, studying primates helps anthropologists comprehend humanity from a biological perspective, which contributes to anthropology\u2019s commitment to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_930\">holism,<\/a><\/strong> the idea that the parts of a system interconnect and interact to make up the whole.<\/p>\n<figure style=\"width: 242px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2023\/05\/image3.png\" alt=\"A person using binoculars to look at monkeys.\" width=\"242\" height=\"354\" \/><figcaption class=\"wp-caption-text\">Figure 7.1: The author observing patas monkeys from a distance in Laikipia, Kenya. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-6\/\">Karin Enstam Jaffe observing patas monkeys in Laikipia, Kenya (Figure 6.5)<\/a> by Rebecca Chancellor is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_932\">Ethology<\/a> <\/strong>is the study of animal behaviour, while <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_934\">primatology <\/a><\/strong>is the study of primate behaviour. People who study primates are called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_936\">primatologists<\/a><\/strong>. Research on primates can be conducted in the field (i.e., on wild primates) or in captivity (i.e., zoos) and may or may not involve experiments, such as playing recorded alarm calls to see how individuals react. Unlike some other Science, Technology, Engineering, and Math (STEM) fields, primatology has a long history of research conducted by women (see \u201cSpecial Topic: Women in Primatology\u201d). Primatologists come from many different disciplines, have diverse backgrounds, and study primates for different reasons. Biologists study primates as examples of evolutionary theories like natural selection, and to understand behaviours as <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_938\">adaptations<\/a><\/strong>, or traits with a function that increases <strong>fitness<\/strong>, i.e. an individual\u2019s survival and\/or reproduction. Primate intelligence is of interest to psychologists who want to learn more about deception or cooperation and to linguists interested in the principles of communication and language. Ecologists consider how primates interact with the habitats they occupy, and conservationists examine how primates are affected by deforestation, poaching, or illegal animal trade (see Appendix B: Primate Conservation for more information on these topics). Biological anthropologists, like myself (Figure 7.1), who study primates are interested in learning about their social complexity, and ecological and behavioural variation, to better understand the biological basis of human behaviour. And, similar to biologists, we also explore how primate behaviour is adaptive and contributes to individual fitness. Like other sciences, primatology is only as strong as its researchers, methods, and theories, and the field has benefitted recently from efforts to increase diversity and reckon with its colonialist past, as discussed below in \u201cSpecial Topic: Women in Primatology.\u201d<\/p>\n<p class=\"import-Normal\">Humans share many traits in common with primates. As you learned in Chapter 5, some of these traits are similar due to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_944\">homology<\/a><\/strong>, traits both species inherited from a common primate ancestor. For example, like most other primates, humans are social animals who live in groups. Group living did not evolve independently in humans and other primates. Rather, group living is a trait that evolved in a primate ancestor, and because it benefited survival, it was retained in the species\u2019 <strong>descendants<\/strong> (or the species that come after the ancestor species). In contrast, humans and other primates can have similar traits that evolved independently, which is called <strong>analogy<\/strong>. For example, both humans and Japanese macaques (<em>Macaca fuscata<\/em>) use natural hot springs (Figures 7.2a &amp; b). Research on these monkeys indicates that sitting in hot springs reduces stress and helps keep them warm, much as it does for humans (Takeshita Et al. 2018). But this behaviour is not the result of humans and Japanese macaques having a shared ancestor who used hot springs. Rather, the behaviour arose independently in two species that both occupy northerly environments and adapted to cold climates using a similar behaviour. Studying the homologous traits we share with other primates, like living in groups, helps us develop hypotheses about human behaviours as adaptations, which in turn helps us develop models for the behaviour of our human ancestors. Studying analogous traits, like hot springs use, allows us to better understand the effects of ecological variables on morphology and behaviour of both primates and humans, living and extinct.<\/p>\n<figure id=\"attachment_191\" aria-describedby=\"caption-attachment-191\" style=\"width: 2161px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-189 size-full\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.2.jpg\" alt=\"Left, a man in a hot spring. Right, monkeys in a hot spring.\" width=\"2161\" height=\"803\" \/><figcaption id=\"caption-attachment-191\" class=\"wp-caption-text\">Figure 7.2a\/b: Both humans (left) and Japanese macaques (right) use natural hot springs to reduce stress and relax. This similar trait arose independently in the two species, making it a good example of analogy. Credit: 7.2a. <a href=\"https:\/\/pixabay.com\/photos\/hot-spring-landscape-man-mountain-1846721\/\">Hot Spring Landscape<\/a> by <a href=\"https:\/\/pixabay.com\/users\/pexels-2286921\/?utm_source=link-attribution&amp;utm_medium=referral&amp;utm_campaign=image&amp;utm_content=1846721\">Pexels<\/a> has been modified (cropped) and 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>. 7.2b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Jigokudani_hotspring_in_Nagano_Japan_001.jpg\">Jigokudani hotspring in Nagano Japan 001<\/a> by Yosemite 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<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Women in Primatology<\/h2>\n<p class=\"import-Normal\">While many STEM fields have traditionally been, and continue to be, dominated by men, primatology has a long history of significant research conducted by women. This is due, in part, to the fact that three of the most well-known primatologists are women. In the early 1960s, British paleoanthropologist Louis Leakey (discussed in Chapters 9 and 10) was looking for students to study the great apes in hopes of shedding light on the behaviours of our early ancestors. He chose Jane Goodall (Figure 7.3a) to study chimpanzees (<em>Pan troglodytes<\/em>), Birute Galdikas (Figure 7.3b) to study Bornean orangutans (<em>Pongo pygmaeus<\/em>), and Dian Fossey (Figure 7.3c) to study mountain gorillas (<em>Gorilla<\/em><em> beringei beringei<\/em>). The work of these three women, sometimes referred to as Leakey\u2019s \u201cTrimates,\u201d has transformed our understanding of ape (and primate) behaviour.<\/p>\n<figure id=\"attachment_190\" aria-describedby=\"caption-attachment-190\" style=\"width: 626px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-190\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.3-1.jpg\" alt=\"Jane Goodall, Birute Galdikas, and Dian Fossey.\" width=\"626\" height=\"209\" \/><figcaption id=\"caption-attachment-190\" class=\"wp-caption-text\">Figure 7.3a-c: Louis Leakey\u2019s \u201cTrimates\u201d (left to right): a. Jane Goodall\u2019s research on the Gombe chimpanzees spans over half a century; b. Birute Galdikas\u2019s research and rescue work on behalf of orangutans spans 40 years; c. Dian Fossey studied mountain gorillas in Rwanda for almost 20 years, until her murder in 1985. Credit: a. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Jane_Goodall_HK.jpg\">Jane Goodall HK<\/a> by Jeekc has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\"> CC BY-SA 3.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Dr_Birute_Galdikas.jpg\">Dr Birute Galdikas<\/a> by<a href=\"https:\/\/www.flickr.com\/photos\/sfupamr\/\"> Simon Fraser University - University Communications<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\"> CC BY 2.0 License<\/a>. c. <a href=\"https:\/\/www.flickr.com\/photos\/mary-lynn\/2925879356\">US-223658 Dian Fossey<\/a> by<a href=\"https:\/\/www.flickr.com\/photos\/mary-lynn\/2925879356\/\"> Mary-Lynn<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\"> CC BY 2.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Arriving at the Gombe Stream Reserve in Tanzania in 1960, Jane Goodall was one of the first scientists to conduct a long-term study of wild nonhuman primates. Before then, most studies lasted less than a year and were often zoo-based. By 1961, she had made two astounding observations that forced us to reconsider what differentiates humans from the rest of the primate order. She observed chimpanzees eating a colobus monkey, the first reported evidence of meat eating in our closest relatives; later observing them hunting and sharing meat. And she discovered that chimpanzees make and use tools by stripping leaves off twigs to \u201cfish\u201d for termites. Her work, spanning several decades, has produced long-term data on chimpanzee mating strategies, mother-infant bonds, and aggression. In the mid-1980s, Goodall transitioned from field researcher to conservationist and activist, advocating for the humane use of nonhuman animals (Stanford 2017).<\/p>\n<p class=\"import-Normal\">Birute Galdikas began her study of orangutans in Kalimantan, Borneo, in 1971. Hers was the first long-term study conducted on the Bornean orangutan. Galdikas and her colleagues have collected over 150,000 hours of observational data, focusing on the life histories of individual orangutans. While conducting behavioural research, Galdikas discovered that the pet trade and habitat loss were adversely affecting the orangutan population. Eventually, Galdikas\u2019s conservation efforts began to extend beyond advocacy and into rehabilitation and forest preservation (Bell 2017). If you would like to learn more about primate conservation efforts, please see Appendix B: Primate Conservation.<\/p>\n<p class=\"import-Normal\">In 1967, Dian Fossey began her long-term study of mountain gorillas and founded the Karisoke Research Centre in Rwanda. Her and her colleagues\u2019 research, over several decades, revealed much about gorilla social behaviour, ecology, and life history. Her efforts also led to the development of mountain gorilla conservation programs. However, she was a controversial figure, as discussed below. Fossey was murdered in December 1985; the case remains unsolved (Stewart 2017).<\/p>\n<h3 class=\"import-Normal\"><strong>Decolonizing Primatology<\/strong><\/h3>\n<p class=\"import-Normal\">Recently, the movement to <strong>decolonize<\/strong> primatology, by understanding and highlighting the theories and research of non-Western individuals and perspectives, has gathered steam. This movement draws attention to the maltreatment of local people by Western primatologists. For example, Michelle Rodrigues (2019) argues that it's time we stop focusing on the scientific and conservation contributions of Dian Fossey and acknowledge that her \"active conservation\" techniques included kidnapping and torturing local Rwandans who were known as, or suspected to be, gorilla poachers. Rodrigues (2019) argues:<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 36pt;text-indent: 0pt\">The image of Fossey, a white American woman, whipping and torturing black African poachers is evocative of the behaviour of white slaveholders in the American South. It is appalling enough to think of that behaviour occurring in the 1850s; there is no way we can explain Fossey\u2019s behaviour in the 1970s as the product of \u201ca different time.\u201d Yet, almost three decades later, the romantic notion of a noble martyr who died for her devotion to gorillas prevails, and these terrifying actions are often described as simply unorthodox methods. Perhaps these truths are softened due to fears that the reality of this legacy would harm gorilla conservation efforts. But memorializing her as a martyr and patron saint of gorilla conservation demands that we forget the cruel acts she advocated for and performed.<\/p>\n<p class=\"import-Normal\">Further, Louis Leakey\u2019s installment of Goodall, Galdikas, and Fossey to study chimpanzees, orangutans, and mountain gorillas, respectively, is itself viewed as recapitulating the colonial legacy in Africa and Asia. Given that Leakey was the offspring of British missionaries, Rodrigues (2019) argues, it is no accident that he was willing to mentor British and American women, while overlooking women from Africa and Asia as potential researchers. This leads us to another level of the decolonizing movement, which aims to highlight the research of non-Western primatologists, particularly those living in what primatologists refer to as \u201chabitat countries\u201d that are home to living primates. As you will see in this chapter, scientists from diverse backgrounds are active contributors to exciting research on primates around the world.<\/p>\n<\/div>\n<h2 class=\"import-Normal\">Ecology<\/h2>\n<p class=\"import-Normal\">The more than 600 species and subspecies of living primates are highly diverse in their dietary preferences and the habitats they occupy. In this section we\u2019ll briefly discuss aspects of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_950\">ecology<\/a><\/strong>, or the relationship between organisms and their physical surroundings, that impact a primate\u2019s life, the foods they eat, and the other species with whom they interact.<\/p>\n<h3 class=\"import-Normal\"><strong>Primate Diets<\/strong><\/h3>\n<p class=\"import-Normal\">Diet may be the most important variable influencing variation in primate morphology, behaviour, and ecology. Most primates are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_952\">omnivores<\/a><\/strong> who ingest a variety of foods in order to obtain appropriate levels of protein, carbohydrates, fats, and fluids, but one type of food often makes up the majority of each species\u2019 diet. You learned about the dental and digestive adaptations of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_954\">frugivores<\/a><\/strong> (who feed primarily on fruit), <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_956\">folivores<\/a><\/strong> (whose diet consists mostly of leaves), and <strong>insectivores <\/strong>(who eat mainly insects) in Chapter 6, so we will not discuss them again here.<\/p>\n<h4 class=\"import-Normal\"><em>Body Size and Diet<\/em><\/h4>\n<figure id=\"attachment_191-2\" aria-describedby=\"caption-attachment-191-2\" style=\"width: 559px\" class=\"wp-caption alignright\"><img class=\"wp-image-191\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.4-1.jpg\" alt=\"A tarsier eats a grasshopper. A gorilla eats leaves.\" width=\"559\" height=\"237\" \/><figcaption id=\"caption-attachment-191-2\" class=\"wp-caption-text\">Figure 7.4a-b: Primates eat different types of food. Small primates, like the spectral tarsier (left), eat mostly insects while large primates, like the mountain gorilla (right), eat mostly leaves. Credit: a. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Spectral_Tarsier_Tarsius_tarsier_(7911549768).jpg\">Spectral Tarsier Tarsius tarsier (7911549768)<\/a> by<a href=\"https:\/\/www.flickr.com\/people\/65695019@N07\"> Bernard DUPONT<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/legalcode\"> CC BY-SA 2.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Mountain_gorilla_(Gorilla_beringei_beringei)_eating.jpg\">Mountain gorilla (Gorilla beringei beringei) eating<\/a> by<a href=\"https:\/\/www.sharpphotography.co.uk\/\"> Charles J Sharp<\/a> (creator QS:P170,Q54800218) has been modified (cropped) 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>Insects are a high-quality food, full of easily digestible protein and high in calories that meet most of a primate\u2019s dietary needs. Although all primates will eat insects if they come upon them, those species that rely most heavily on insects tend to be the smallest. Why? Because larger primates simply cannot capture and consume enough insects every day to survive. Because of their small size (less than 150 g), spectral tarsiers (<em>Tarsius spectrum<\/em>) have a fast <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_960\">metabolism<\/a><\/strong>, which means they turn food to energy quickly, but they do not need to consume large amounts of food each day. It does not matter to a spectral tarsier that a grasshopper only weighs 300 mg, because the tarsier (<em>Tarsius<\/em>) itself is so small that one grasshopper is a good-size meal (Figure 7.4a). That same grasshopper is not even a snack for an adult male mountain gorilla (<em>Gorilla beringei beringei<\/em>), who may weigh up to 200 kg. Fortunately for gorillas (<em>Gorilla)<\/em>, their large body size means they have a slow metabolism, converting food into energy much more slowly, so they can eat lower quality food that takes longer to digest, provided there is a lot of it. For gorillas, leaves, which are hard to digest but plentiful, fit the bill (Figure 7.4b). Most medium-sized primates are highly frugivorous, and supplement their fruit based diet in ways that correspond with their size: Smaller frugivores tend to supplement with insects, while larger frugivores tend to supplement with leaves.<\/p>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<h4 class=\"import-Normal\"><em>Food Abundance and Distribution<\/em><\/h4>\n<p class=\"import-Normal\">Nutrients are not the only dietary considerations primates must make. They must also ensure that they consume more calories than they use. The abundance and distribution of food affect energy expenditure and calorie intake because they determine how far animals must travel in search of food and how much they must compete to obtain it. <strong>Abundance <\/strong>refers to how much food is available in a given area while <strong>distribution<\/strong> refers to how food is spread out. In terms of abundance, food is either plentiful or scarce (Figure 7.5a\u2013b). Food is distributed in one of three ways: uniformly (Figure 7.6a), in clumps (Figure 7.6b), or randomly (Figure 7.6c). In general, higher-quality foods, like fruit and insects, are less abundant and have patchier distributions than lower-quality foods, like leaves. Primates who eat fruit or insects usually have to travel farther to find food and burn more calories in the process. Abundance and distribution of food is another reason why larger primates tend to rely more heavily on leaves than either fruit or insects.<\/p>\n<figure style=\"width: 619px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image2-2.png\" alt=\"Two squares with different amounts of dots. \" width=\"619\" height=\"286\" \/><figcaption class=\"wp-caption-text\">Figure 7.5a-b: Two types of food abundance. Food is plentiful when there is a lot of it in a given area (left). Food is scarce when there is not very much of it in a given area (right). Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-6\/\">Food abundance and food scarcity (Figure 6.7)<\/a> by Karin Enstam Jaffe 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<figure style=\"width: 690px\" class=\"wp-caption aligncenter\"><img style=\"font-size: 1em\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-2.png\" alt=\"Three squares with dots in different formations. \" width=\"690\" height=\"214\" \/><figcaption class=\"wp-caption-text\">Figure 7.6a-c: Three types of food distribution. a. Food has a uniform distribution when it is spread out evenly in the environment. b. Food has a clumped distribution when it is found in patches. c. Food is randomly distributed when it has neither uniform nor clumped distribution. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-6\/\">Food distribution patterns (Figure 6.8)<\/a> by Karin Enstam Jaffe 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<h3 class=\"import-Normal\"><strong>Community Ecology<\/strong><\/h3>\n<p class=\"import-Normal\">Primates are members of broader ecological communities composed of other species, including other primates, predators, parasites, and even humans. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_966\">Community ecology<\/a><\/strong> deals with the relationships and interactions between different organisms that occupy the same habitat. Interactions with <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_968\">conspecifics<\/a> <\/strong>(members of the same species) and <strong>heterospecifics<\/strong> (members of different species) are critical aspects of ecological communities. Some habitats support highly diverse <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_972\">primate communities<\/a><\/strong> consisting of 10 or more primate species. How can so many primate species occupy the same area and avoid competition? In most cases, the primate species that live together occupy different <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_974\">niches<\/a><\/strong>, which means they do not meet their needs for food and shelter in the exact same way. Two species can avoid competition by eating different kinds of food, living at different levels of a forest, or even searching for food at different times of day. Because tropical rainforests, like Manu National Park in Peru, are highly variable, with many habitats and many sources of food and shelter, there are many different niches for multiple species to exploit, and large primate communities can result (Figure 7.7).<\/p>\n<figure style=\"width: 710px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5-1-1.png\" alt=\"Eight primate species.\" width=\"710\" height=\"550\" \/><figcaption class=\"wp-caption-text\">Figure 7.7: Eight of the 14 primate species in Manu National Park, Peru. Top row, left to right: Goeldi\u2019s marmoset (Callimico goeldi), Rio Tapaj\u00f3s saki (Pithecia irrorata), tufted capuchin (Sapajus apella); middle row, left to right: emperor tamarin (Saguinus imperator), black-headed night monkey (Aotus nigriceps), Bolivian red howler (Alouatta sara); bottom row, left to right: black-capped squirrel monkey (Saimiri boliviensis), Peruvian spider monkey (Ateles chamek). Credit: Primate species in Manu National Park original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Jaffe 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:\/\/www.flickr.com\/photos\/31223088@N08\/5582747190\/\">Tamarin Baby\/Goeldi\u2019s Monkey<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/31223088@N08\/\">stefan_fotos<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\">CC BY 2.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Pithecia_irrorata_-Brazil-8b.jpg\">Pithecia irrorata -Brazil-8b<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/9092428@N04\">Ana_Cotta<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/deed.en\">CC BY 2.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Tufted_capuchin_on_a_branch_in_Singapore.jpg\">Tufted Capuchin on a Branch in Singapor<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Basile_Morin\">Basile_Morin<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Tamarin_portrait.JPG\">Tamarin Portrait<\/a> by <a href=\"https:\/\/sites.google.com\/site\/thebrockeninglory\/?pli=1\">Brocken Inaglory<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/deed.en\">CC BY-SA 3.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Aotus_nigriceps_1.jpg\">Aotus nigriceps 1<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/dusantos_bh\/\">DuSantos<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/deed.en\">CC BY 2.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Alouatta_sara_%28Bolivian_red_howler%29.jpg\">Aloutta sara (Bolivian Red Howler)<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/nacho_dayz\/\">Raul Ignacio<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\">CC BY-SA 2.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Black-capped_squirrel_monkey_%28Chalalan%29.jpg\">Black-Capped Squirrel Monkey (Chalalan)<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Rodrigo_Mariaca\">Rodrigo Mariaca<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0 License<\/a>; <a href=\"https:\/\/www.flickr.com\/photos\/eye1\/3185562151\/\">Maquisapa (Spider Monkey)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/eye1\/\">Ivan Mlinaric<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\">CC BY 2.0 License<\/a>].<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Competitive Interactions<\/em><\/h4>\n<p class=\"import-Normal\">Although species living in the same location often occupy different niches to avoid competition, when a resource that is important for survival or reproduction is scarce, individuals will compete to obtain that resource. This is a central tenet of Charles Darwin\u2019s theory of evolution by natural selection (see Chapter 2 &amp; 3). Competition between primates takes two forms: Individuals engage in <strong>direct competition<\/strong>, which involves physical interaction between individuals (such as fighting), over resources that are large and worth defending (fruit is a good example of a food resource over which primates will fight). Individuals engage in <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_978\">indirect competition<\/a><\/strong>, in which there is no physical interaction between individuals, when a resource is small. Primates often engage in indirect competition for insects, like grasshoppers, that are eaten quickly, often before another individual arrives on the scene. Primates may engage in direct and\/or indirect competition with members of their own group, with members of other groups of conspecifics, or with heterospecifics.<\/p>\n<h4 class=\"import-Normal\"><em>Predator-Prey Interactions<\/em><\/h4>\n<p class=\"import-Normal\">The plants and animals that primates eat are an important part of their ecological community. In addition to insects, many primates incorporate some <strong>vertebrate<\/strong> (animals with an internal spinal column or backbone) prey into their diet. Often, predation by primates is opportunistic, occurring because the prey happens to be in the right place at the right time. I\u2019ve observed vervets (<em>Chlorocebus pygerythrus<\/em>) opportunistically killing lizards by smashing them against a rock or tree trunk and eating them. More rarely, hunting is deliberate and cooperative. In some chimpanzee (<em>Pan troglodytes<\/em>) populations, hunts involve multiple individuals, each of whom plays a specific role and is rewarded afterward with a share of the prey that has been captured (Samuni Et al. 2018).<\/p>\n<p class=\"import-Normal\">All primates are susceptible to predation by mammalian <strong>carnivores <\/strong>(animals whose diet consists primarily of animal tissue (e.g., Figure 7.8a), reptiles (e.g., Figure 7.8b), or birds of prey (e.g., Figure 7.8c). Although the specific predators found in an ecological community differ based on geography, smaller primates always fall prey to a wider range of predators. Because predators are diverse in their hunting tactics, primates have evolved a wide range of tactics to avoid or escape them. We will discuss some of these behavioural adaptations later in this chapter in the section titled \u201cWhy Do Primates Live in Groups?.\u201d<\/p>\n<p>&nbsp;<\/p>\n<\/div>\n<figure id=\"attachment_210\" aria-describedby=\"caption-attachment-210\" style=\"width: 765px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-195\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.8-scaled-1.jpg\" alt=\"A leopard, python, and harpy eagle.\" width=\"765\" height=\"229\" \/><figcaption id=\"caption-attachment-210\" class=\"wp-caption-text\">Figure 7.8a-c: Examples of primate predators: the Indian leopard (Panthera fusca) is an example of a mammalian carnivore (top left), the South African python (Python natalensis) is an example of a reptilian predator (bottom left), and the harpy eagle (Harpia harpyja) of Central and South America is an example of a bird of prey (right). Credit: a. <a href=\"https:\/\/www.flickr.com\/photos\/srikaanth-sekar\/9814267145\/\">Leopard<\/a> by<a href=\"https:\/\/www.flickr.com\/people\/srikaanth-sekar\/\"> Srikaanth Sekar<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/\"> CC BY-SA 2.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Python_natalensis_G._J._Alexander.JPG\">Python natalensis G. J. Alexander<\/a> by Graham J. Alexander, University of the Witwatersrand, USGS, is in the<a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\"> public domain<\/a>. c. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Harpy_Eagle_clutching_captured_bird_-_Itirapina_Reserve.jpg\">Harpy Eagle clutching captured bird - Itirapina Reserve<\/a> by Jonathan Wilkins has been modified (cropped) and 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<div class=\"__UNKNOWN__\">\n<h4 class=\"import-Normal\"><em>Mutualistic Interactions<\/em><\/h4>\n<p class=\"import-Normal\">So far, we've discussed competitive and predator-prey interactions in primate communities. But there are some interactions (between different primate species and between primates and other species) that are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_984\">mutualistic<\/a><\/strong>, which is when organisms of different species work together, each benefiting from the interaction or relationship. One example is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_986\">seed dispersal<\/a><\/strong>, which is the process by which seeds move away from the plant that produced them in preparation for germination and becoming a new plant. When seeds are dispersed by animals, like primates, it is an example of mutualism. The primate eats the fruit of a plant, which provides nutrients for its body, and in the process ingests the plant\u2019s seeds. Later, it deposits the seeds at another location as a pile of fertilizer.<\/p>\n<p class=\"import-Normal\">Another example of mutualism is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_988\">polyspecific associations<\/a><\/strong>, which are associations between two or more different species that are maintained by behavioural changes by at least one of the species. While some associations are short in duration, others are semi-permanent. The mutualistic benefits of polyspecific associations include one species gaining access to food that would otherwise have been inaccessible or being alerted to the presence of predators that they would not have not have known were present otherwise. In some cases, individuals seem to recognize and seek out specific members of another species. Twenty years of observations on chimpanzees and Western lowland gorillas (<em>Gorilla gorilla gorilla<\/em>) in the Republic of Congo has revealed social ties (some might call them friendships) between individual chimpanzees and gorillas that last for years and occur in a variety of social contexts, including play (Sanz Et al. 2022).<\/p>\n<h4 class=\"import-Normal\"><em>Parasite-Host Interactions<\/em><\/h4>\n<p class=\"import-Normal\">Primates are hosts for a variety of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_990\">parasites<\/a><\/strong>, which are organisms that live in or on another organism (the host). Parasites come in many forms and pose varying levels of danger to the host. Blood parasites cause diseases like yellow fever and malaria. Skin parasites include fleas and ticks, which feed on the host\u2019s blood, and botflies, which lay eggs in the host\u2019s flesh. Bot fly larvae feed on the host\u2019s flesh as they develop and eventually (if not removed) break through the skin at maturity. Gut parasites, like tapeworms, get into the intestines and feed off of the food that is being digested by the host. Because most primates live in groups (see the \u201cPrimate Societies\u201d section of this chapter), the tendency for <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_992\">social transmission<\/a><\/strong> of parasites, or the transfer of parasites from one individual to another, is high. Primates have evolved mechanisms to avoid parasite infection, including switching sleeping and feeding sites so as to avoid parasites. Mandrills (<em>Mandrillus sphinx<\/em>) have been shown to avoid grooming infected conspecifics as well as to avoid their feces, which smell different than the feces of individuals who are not infected with parasites (Poirotte Et al. 2017). Other primates, including chimpanzees, appear to self-medicate when infected with parasites by ingesting plants that have antiparasitic properties (Krief Et al. 2005).<\/p>\n<h4 class=\"import-Normal\"><em>Human-Primate Interactions<\/em><\/h4>\n<p class=\"import-Normal\">Humans are part of many primate communities and our relationship with our closest relatives is often complicated. In some areas, humans hunt primates for their meat or as trophies, or so they can sell the infants as pets. As the human population increases in size, our demand for natural resources, like wood to build houses or land on which to grow food, also increases, often at the expense of pristine primate (and other animal) habitat. As their natural habitat shrinks, primates search for food in areas occupied by humans and may be shot as crop-raiding pests. While deforestation, hunting, and the pet trade are examples of ways in which humans negatively affect the lives of primates, some human-primate interactions are beneficial. In some parts of the world primates are central to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_994\">ecotourism<\/a><\/strong>, which focuses on nature-based attractions to educate tourists and uses economically and ecologically sustainable practices. Perhaps one of the greatest success stories of ecotourism involves the mountain gorillas of Rwanda (see Figure 7.4b). After internal conflict plagued Rwanda during the 1990s, the Virunga Mountains area developed gorilla-based tourism to aid in socioeconomic development and to bring stability to the region. This process not only helped to increase mountain gorilla populations but was also able to generate enough income to cover the operation costs of three national parks and provide income and other benefits to people living in the area (Maekawa Et al. 2013). You can learn more about human-primate interactions in Appendix B: Primate Conservation.<\/p>\n<h2 class=\"import-Normal\">Primate Societies<\/h2>\n<p class=\"import-Normal\">Unlike many other animals, primates are highly social and many live in stable groups consisting of adult males and females, even outside the<strong> <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_996\">breeding season<\/a><\/strong>, when females are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_998\">receptive<\/a><\/strong> and available for mating because they are not pregnant or nursing. Indeed, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1000\">sociality<\/a><\/strong>, or the tendency to form social groups, is a key behavioural adaptation of the order primates (see Chapter 6). This has led primatologists to ask two questions: \u201cWhy do primates live in groups?\u201d and \u201cWhat types of groups do primates live in?\u201d<\/p>\n<h3 class=\"import-Normal\"><strong>Why Do Primates Live in Groups?<\/strong><\/h3>\n<p class=\"import-Normal\">Primates live in groups when the benefits of doing so exceed the costs. Although there are many potential benefits to group living, enhanced feeding competition and predator avoidance are important benefits for many group living primates. When primates feed on high-quality, scarce food (like fruit), larger groups are more successful in competition with other groups. For example, in a long-term study of vervets in Kenya\u2019s Amboseli National Park, larger vervet groups had larger and better <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1002\">home ranges<\/a><\/strong>, which is the area in which a group regularly moves around as it performs its daily activities, including searching for food and water. Females in larger groups had higher average infant and female survival rates than the smallest group. Because pregnancy and nursing are energetically expensive for females, female <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1004\">reproductive success<\/a><\/strong>, or genetic contribution to future generations (measured by the number of offspring produced), is limited by access to food. Although living in a group means females compete with members of their own group for food, the benefits of being a member of a larger vervet group outweigh the costs (Cheney &amp; Seyfarth 1987).<\/p>\n<p class=\"import-Normal\">However, because they contain more individuals, larger groups are more likely to attract the attention of predators compared to smaller groups. This is one of the reasons that primates who rely on <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1006\">crypsis<\/a><\/strong>, or the ability to avoid detection by others, including predators, are often <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1008\">solitary<\/a> <\/strong>(the term used to describe individuals who do not live together with other members of their species) and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1010\">nocturnal<\/a><\/strong>, or active at night. If an animal is already hard to see because it is active at night, then moving quietly in small groups is a good strategy to avoid detection by predators. The slow loris (<em>Nycticebus coucang<\/em>) of Southeast Asia is a good example of this strategy (Figure 7.9a). Nocturnal and solitary, the slow loris moves slowly and quietly as its primary strategy to avoid detection (Wiens &amp; Zitzmann 2003). In contrast, primates who live in large groups and are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1012\">diurnal<\/a><\/strong>, or active during the day (like gelada baboons [<em>Theropithecus gelada<\/em>]; Figure 7.9b) cannot avoid detection by predators. Instead, group-living primates rely on behaviours that alert others to the presence of danger and\/or deter predators, including shared <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1014\">vigilance<\/a><\/strong> (watchful behaviour to detect potential danger), <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1016\">mobbing<\/a><\/strong> (the act of cooperatively attacking or harassing a predator), and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1018\">alarm calling<\/a><\/strong> (vocalizations emitted by social animals in response to danger). We will discuss alarm calls in the <em>Communication<\/em> section.<\/p>\n<figure id=\"attachment_196\" aria-describedby=\"caption-attachment-196\" style=\"width: 761px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-196\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.9.jpg\" alt=\"A slow loris. A group of gelada baboons.\" width=\"761\" height=\"269\" \/><figcaption id=\"caption-attachment-196\" class=\"wp-caption-text\">Figure 7.9a\/b: Some primates, like the slow loris (left), are solitary and spend most of their time alone. However, most primates, like the gelada baboon (right), live in groups of varying sizes. Credit: a. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Slow_Loris.jpg\">Slow Loris<\/a> by Jmiksanek is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\"> CC BY-SA 3.0 License<\/a>. b. <a href=\"https:\/\/www.flickr.com\/photos\/39997856@N03\/7588490544\">Field of baboons<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/39997856@N03\/\">mariusz kluzniak<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/2.0\/\">CC BY-NC-ND 2.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>What Types of Groups Do Primates Live In? <\/strong><\/h3>\n<p class=\"import-Normal\">Primates vary with regard to the types of groups in which they live. A <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1020\">social system<\/a> <\/strong>describes a set of social interactions and behaviours that is typical for a species. The components that make up a species\u2019 social system include:<\/p>\n<ul>\n<li class=\"import-Normal\">Group size, which refers to the number of individuals that typically live together. Primate group size can be highly variable, ranging from one or a few individuals, to a few dozen, upward to several hundred individuals.<\/li>\n<li class=\"import-Normal\">Group composition describes group membership in terms of age class (e.g., adult, juvenile, infant) and sex. In some primates, groups consist of a mother and her dependent offspring while in others, one adult male lives long-term with one adult female and their dependent offspring. In other species, one or more adult males live with multiple females and their offspring.<\/li>\n<li class=\"import-Normal\">A species\u2019 <strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1022\">mating system<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"> refers to which male(s) and female(s) mate. The terms that describe a mating system (e.g., <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1024\">polygyny<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">, in which one male mates with multiple females) are sometimes used to describe a primate species\u2019 social system, but a mating system is one component of the species\u2019 social system. For example, two species might both have polygynous mating systems, but in one species, the group is composed of one male and multiple females, while members of the other species live as solitary individuals.<\/span><\/li>\n<li class=\"import-Normal\"><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1026\">Ranging behaviour<\/a> <\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">refers to the way in which animals move about their environment. Most primate species have a home range, where they perform their daily activities. Some primates defend a <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1028\">territory<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"> which is the part of the home range that the group actively guards in an attempt to keep out conspecifics.<\/span><\/li>\n<li class=\"import-Normal\"><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1030\">Dispersal<\/a> <\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">patterns describe which sex moves to a new group to reproduce. In most primate species, males disperse because the benefits of dispersal, including increased access to mates and reduced competition from other males, outweigh the costs of migrating into a new group, which often comes with aggression from current group members. For many female primates, the opposite is true: females usually benefit from remaining <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1032\">philopatric<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">, or in the group of their birth. This allows them to maintain strong alliances with female relatives, which helps them compete successfully against other groups for food. In solitary species, offspring of both sexes leave their mother\u2019s home range and become solitary. If this did not happen, the species would not be solitary. Even though both sexes disperse in solitary species, males usually disperse farther than females.<\/span><\/li>\n<li class=\"import-Normal\">Social interactions describe the ways in which individuals interact with members of their own and other groups of conspecifics. <strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1034\">Affiliative<\/a> <\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">(i.e., friendly or nonaggressive) behaviours include <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1036\">grooming<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"> (picking through the fur of another individual), playing, or <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1038\">coalitions<\/a> <\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">(temporary alliances between individuals). <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1040\">Agonistic<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"> (i.e., aggressive) behaviours include fighting over food or fighting over access to mates. In groups that contain multiple adult individuals of the same sex, it is common to have a <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1042\">dominance hierarchy<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">, or a group of individuals that can be ranked according to their relative amount of power over others in the hierarchy. Initially, dominance hierarchies are established through the outcome of conflicts. Individuals who lose conflicts with others are <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1044\">subordinate<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"> (or low rank) to those who win them. Those who win conflicts are <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_744\">dominant<\/a> <\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">(or high rank). Dominant individuals gain access to resources, like food or mates, before subordinates. Once a hierarchy is established, agonism decreases because everyone \u201cknows their place.\u201d<\/span><\/li>\n<\/ul>\n<p class=\"import-Normal\">The main types of primate social systems are as follows: solitary; single-male, single-female; single-male, multi-female; multi-male, multi-female; fission-fusion; and multi-male, single-female. These types are discussed below.<\/p>\n<h4 class=\"import-Normal\"><em>Solitary<\/em><\/h4>\n<p class=\"import-Normal\">Recall that the term <em>solitary<\/em> is used to describe species in which individuals do not live or travel together with other members of the same species, except for mothers and unweaned offspring. Males typically occupy a large home range or territory that overlaps the home ranges of multiple females, with whom they mate (Figure 7.10a). Because one male mates with multiple females, the mating system of solitary primates is polygyny. Social interactions between adults are limited but because some males do not get to mate, competition between males is intense. When males compete physically, they benefit from large body size and weaponry. The result is<strong> <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1046\">sexual dimorphism<\/a><\/strong>, when males and females look different from one another. Both males and females disperse, although males move farther from their mother than females. The nocturnal West African potto (<em>Perodicticus <\/em><em>potto<\/em>; Figure 7.10b) is solitary. Bornean orangutans, which are diurnal, are also solitary.<\/p>\n<\/div>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_210-2\" aria-describedby=\"caption-attachment-210-2\" style=\"width: 620px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-197\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.10-01-1.jpg\" alt=\"Grouping pattern description is available in caption. A potto in a tree at night also shown.\" width=\"620\" height=\"272\" \/><figcaption id=\"caption-attachment-210-2\" class=\"wp-caption-text\">Figure 7.10a-b: Illustration of a solitary species\u2019 grouping pattern is shown on the left. Key: square = adult male; dot = adult female; open circle represents the outline of the male\u2019s home range; open oval represents individual female home ranges. The West African potto is a solitary primate (right). <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. Polygyny in a Solitary Social System original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Enstam Jaffe is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>. b. <a href=\"https:\/\/www.flickr.com\/photos\/nikborrow\/31307385633\">West African Potto Perodicticus potto Kakum National Park, Ghana<\/a> by<a href=\"https:\/\/www.flickr.com\/people\/nikborrow\/\"> Nik Barrow<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/2.0\/\"> CC BY-NC 2.0 License<\/a>.<\/figcaption><\/figure>\n<div class=\"__UNKNOWN__\">\n<h4 class=\"import-Normal\"><em>Single-Male, Single-Female<\/em><\/h4>\n<p class=\"import-Normal\">Primate species in which an adult male and adult female live together with their dependent offspring have a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1048\">single-male, single-female<\/a><\/strong> social system, sometimes referred to as a \u201cfamily,\u201d with group sizes between two and five individuals. The adult male and adult female engage in behaviours that strengthen their social relationship, or <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1050\">pair bond<\/a><\/strong>, including mutual grooming and resting together. The pair defend a territory (Figure 7.11a) and keep same-sex individuals away from their mate. The adult male and adult female mate with each other, so the mating system is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1052\">monogamy<\/a><\/strong>, although mating outside the pair bond may occur. Species with monogamous mating systems are usually <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1054\">sexually monomorphic<\/a><\/strong> (males and females look similar) because competition for mates is relaxed since most males are able to obtain a mate. Males are usually confident that they are the father of their mate\u2019s infant, so they help with offspring care by carrying the infant when it is not nursing. Once offspring are sexually mature, both males and females disperse. As with solitary species, males disperse farther from their parents than females. Bolivian Gray titi monkeys (<em>Plecturocebus donacophilus<\/em>) are an example of a species that has a single-male, single-female social system. One of their signature behaviours is tail twining, when two individuals sit with their tails wrapped around each other (Figure 7.11b). This behaviour reinforces the social bond among family members and is especially common between the adult male and female. Gibbons (<em>Hylobates<\/em>) and owl monkeys (<em>Aotus<\/em>) also live in single-male, single-female groups.<\/p>\n<figure id=\"attachment_198\" aria-describedby=\"caption-attachment-198\" style=\"width: 608px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-198\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.11-01.jpg\" alt=\"Left: Circle contains one dot (female) and one square (male). Right: Two titi monkeys.\" width=\"608\" height=\"357\" \/><figcaption id=\"caption-attachment-198\" class=\"wp-caption-text\">Figure 7.11a-b: Illustration of a single-male, single-female grouping pattern is shown on the left. Key: square = adult male; dot = adult female; open circle represents the outline of the group\u2019s territory, which the bonded pair defend against conspecifics. The titi monkey (right) is an example of a primate species with a single-male, single-female social system. Credit: a. Single-Male, Single-Female Social System original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Enstam Jaffe is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Callicebus-brunneus-London-Zoo.jpg\">Two Red Titi Monkeys (Callicebus cupreus) sitting together with their tails intertwined at the London Zoo<\/a> by<a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Stevenj\"> Steven G. Johnson<\/a> has been modified (cropped) and 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<\/div>\n<div class=\"__UNKNOWN__\">\n<h4 class=\"import-Normal\"><em>Single-Male, Multi-Female<\/em><\/h4>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1056\">Single-male, multi-female<\/a><\/strong> groups consist of one adult male living with multiple adult females and their dependent offspring (Figure 7.12a ) . These groups can range from as few as five or ten individuals to as many as 50. Female social relationships are governed by the female dominance hierarchy. Females are usually philopatric and males disperse. Males who are unable to join a group of females may join a bachelor group with other males. Because a single male mates with multiple females, the mating system is polygyny. Species that form single-male, multi-female groups may or may not defend a territory, but the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1058\">resident male<\/a><\/strong>, who lives with a group of females, is aggressive toward other males, who may try to take over the group and become the new resident male. Competition between males to be the resident male of a group is intense, and these species usually display sexual dimorphism, with males being larger than females and possessing large canines. Hanuman langurs (<em>Semnopithecus entellus<\/em>) of India form single-male, multi-female groups (Figure 7.12b). When a new male takes over a group of females and ousts the former resident male, he may commit <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1060\">infanticide<\/a>, <\/strong>or kill the unweaned infants. This is especially likely if the new resident male has not yet mated with any of the females and thus cannot be the infants\u2019 father. This causes the females, who were nursing, to become sexually receptive sooner, increasing the new resident male\u2019s chances of producing offspring (Sharma, Ram, and \u200b\u200bRaipurohit 2010). Gorillas, patas monkeys, and golden snub-nosed monkeys (<em>Rhinopithecus roxellana<\/em>) also live in single-male, multi-female groups.<\/p>\n<p>&nbsp;<\/p>\n<\/div>\n<figure id=\"attachment_210-3\" aria-describedby=\"caption-attachment-210-3\" style=\"width: 749px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-199\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.12.jpg\" alt=\"Left: Circle contains nine dots and one square; outside are three squares. Right: Adult langur and infant.\" width=\"749\" height=\"295\" \/><figcaption id=\"caption-attachment-210-3\" class=\"wp-caption-text\">Figure 7.12a-b: An illustration of the one-male, multi-female grouping pattern is shown on the left. Key: square = adult male; dot = adult female; open circle represents the outline of the group\u2019s home range (or territory). The Hanuman langur (right) is an example of a species with a one-male, multi-female social system. Credit: a. Single-Male, Multi-Female Social System original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Enstam Jaffe is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>. b. <a href=\"https:\/\/www.pexels.com\/photo\/close-up-photo-of-two-gray-langurs-8642891\/\">Close-up of Two Grey Langurs<\/a> by<a href=\"https:\/\/www.pexels.com\/@amitrai10\/\"> Amit Rai<\/a> has been modified (cropped) and is<a href=\"https:\/\/www.pexels.com\/license\/\"> free to use via Pexels<\/a>.<\/figcaption><\/figure>\n<div class=\"__UNKNOWN__\">\n<h4 class=\"import-Normal\"><em>Multi-Male, Multi-Female<\/em><\/h4>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1062\">Multi-male, multi-female<\/a><\/strong> groups consist of multiple adult males living with multiple adult females and their dependent offspring. Although there is more than one adult male, there are more adult females than adult males in the group (Figure 7.13a). Multi-male, multi-female groups can range in size from about ten to as many as 500 individuals. They occupy a home range but may or may not defend a territory. In groups that contain multiple males and multiple females, it is not possible for one male to monopolize all the matings, so the mating system is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1064\">polygamy<\/a><\/strong>, in which multiple males mate with multiple females. However, this does not mean that all males have an equal opportunity to mate with all females. In multi-male, multi-female groups, both males and females form a dominance hierarchy. The male dominance hierarchy determines their access to females for mating in much the same way that a female dominance hierarchy determines a female\u2019s access to food. Because their place in the hierarchy can affect their reproductive success, males compete with each other, but because it is rare for males to be excluded from mating altogether, the level of competition and degree of sexual dimorphism are less extreme than what we see in polygynous species. Usually, females are philopatric and males disperse. Vervet monkeys (Figure 7.13b), ring-tailed lemurs (<em>Lemur catta<\/em>), white-faced capuchins (<em>Cebus capucinus<\/em>), and black-capped squirrel monkeys (<em>Saimiri boliviensis<\/em>) live in multi-male, multi-female groups.<\/p>\n<\/div>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_210-4\" aria-describedby=\"caption-attachment-210-4\" style=\"width: 585px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-200\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.13.jpg\" alt=\"Circle contains twelve dots and three squares. Right: Two vervet monkeys.\" width=\"585\" height=\"267\" \/><figcaption id=\"caption-attachment-210-4\" class=\"wp-caption-text\">Figure 7.13a-b: An illustration of the multi-male, multi-female grouping pattern is shown on the left. Key: square = adult male; dot = adult female; open circle represents the outline of the group\u2019s home range (or territory). Vervet monkeys (right) are an example of a species that lives in multi-male, multi-female groups. Credit: a. Multi-Male, Multi-Female Social System original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Enstam Jaffe is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>. b. <a href=\"https:\/\/www.flickr.com\/photos\/berniedup\/6011902081\/\">Vervet Monkeys (Chlorocebus pygerythrus)<\/a> by<a href=\"https:\/\/www.flickr.com\/people\/65695019@N07\"> Bernard DUPONT<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/deed.en\"> CC BY-SA 2.0 License<\/a>.<\/figcaption><\/figure>\n<div class=\"__UNKNOWN__\">\n<h4 class=\"import-Normal\"><em>Fission-Fusion<\/em><\/h4>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1066\">Fission-fusion<\/a><\/strong> is a fluid social system in which the size and composition of the social group changes, with groups splitting (fission) or merging (fusion) depending on food availability (Pinacho-Guendulain &amp; Ramos-Fern\u00e1ndez 2017). When key resources are scarce, individuals spread out (fission) and move and feed individually or in small subgroups (Figure 7.14a). When key food resources are plentiful, individuals come together (fusion) and individuals travel and feed as a more cohesive group (Figure 7.14a). Fission-fusion social structure is believed to reduce feeding competition when resources are scarce. Because group composition changes over time, species with fission-fusion social systems are referred to as a community. Communities consist of multiple adult males, multiple adult females, and offspring, and group size varies but typically ranges from ten to a few dozen individuals. Females typically disperse and males are philopatric. Thus, community males are related and display unusual forms of cooperation. The mating system associated with fission-fusion is polygamy. Because males are not excluded from mating, competition for mates is relaxed and sexual dimorphism is moderate (males are slightly larger than females). Geoffroy\u2019s spider monkeys (<em>Ateles geoffroyi<\/em>) (Figure 7.14b) and chimpanzees both have fission-fusion social system.<\/p>\n<\/div>\n<figure id=\"attachment_210-5\" aria-describedby=\"caption-attachment-210-5\" style=\"width: 720px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-201\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.14.jpg\" alt=\"Diagrams show fission and fusion. Three spider monkeys.\" width=\"720\" height=\"289\" \/><figcaption id=\"caption-attachment-210-5\" class=\"wp-caption-text\">Figure 7.14a-b: An illustration of the fission-fusion grouping pattern appears on the left. The left illustration represents fission, when females travel and feed independently in individual home ranges within the community boundary. The right illustration represents fusion, when community members form a cohesive group. <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>. Key: square = adult male; dot = adult female; open circle represents the outline of the community boundary. Open ovals represent individual female home ranges when the group fissions. Credits: a. Fission-Fusion Social System original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Enstam Jaffe is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>. b. <a href=\"https:\/\/www.flickr.com\/photos\/berniedup\/49562895393\">Geoffry\u2019s Spider Monkeys (Ateles geoffroyi)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/65695019@N07\">Bernard DUPONT<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/deed.en\"> CC BY-SA 2.0 License<\/a>.<\/figcaption><\/figure>\n<div class=\"__UNKNOWN__\">\n<h4 class=\"import-Normal\"><em>Multi-Male, Single-Female<\/em><\/h4>\n<p class=\"import-Normal\">In <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1068\">multi-male, single-female<\/a> <\/strong>groups, two or more males live with one breeding female, her dependent offspring, and non-breeding females (Figure 7.15a). This type of social system is found in the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1070\">callitrichids<\/a><\/strong>, the primate family that includes marmosets (<em>Callithrix<\/em>; Figure 7.15b) and tamarins (<em>Saguinus<\/em>) of Central and South America. Their groups rarely exceed 15 individuals, and each group actively defends their territory from conspecifics. Although more than one adult female may live in the group, the mating system is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1072\">polyandry<\/a><\/strong> because there is only one breeding female who mates with all of the adult males. This is achieved through <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1074\">reproductive suppression<\/a><\/strong>, which involves the breeding female preventing other females from reproducing through physiological and\/or behavioural means (Digby, Ferrari, &amp; Salzman 2011). This limits the opportunities for other females in the group to become pregnant. Instead, these females, and the males in the group, help raise the breeding female\u2019s offspring. This is referred to as <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1076\">cooperative breeding<\/a><\/strong> and usually takes the form of carrying infants, grooming them, and protecting them from danger (de Oliveira Terceiro &amp; Burkart 2019). Because reproductive opportunities for female tamarins and marmosets are limited, they are very competitive, and females are slightly larger than males, which helps them compete for the breeding spot in a group.<\/p>\n<p>&nbsp;<\/p>\n<\/div>\n<figure id=\"attachment_210-6\" aria-describedby=\"caption-attachment-210-6\" style=\"width: 578px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-202\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.15.jpg\" alt=\"Circle contains two squares, two unmarked dots, and a B dot. Right: Marmosets with twins.\" width=\"578\" height=\"269\" \/><figcaption id=\"caption-attachment-210-6\" class=\"wp-caption-text\">Figure 7.15a-b: An illustration of multi-male, single-female grouping pattern appears on the left. Key: square = adult male; B dot = breeding female; unmarked dot = non-breeding female; open circle represents the outline of the group\u2019s territory, which is defended against conspecifics. The common marmoset (Callithrix jacchus) is an example of a primate species that has this type of social system (right). Credit: a. Multi-Male, Single-Female Social System original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Enstam Jaffe is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Family_of_Common_Marmoset_-_REGUA_-_Brazil_MG_9480_(12930855765).jpg\">Family of Common Marmoset - REGUA - Brazil MG 9480 (12930855765)<\/a> by<a href=\"https:\/\/www.flickr.com\/people\/30818542@N04\"> Francesco Veronesi<\/a> has been modified (cropped) 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<div class=\"__UNKNOWN__\">\n<h2 class=\"import-Normal\">Reproductive Strategies<\/h2>\n<p class=\"import-Normal\">Reproductive strategies have evolved to maximize individual reproductive success. These strategies can be divided into those that deal with offspring production and care (parental investment) and those that maximize mating success (sexual selection). Because the reproductive physiology of male and female primates differs, males and females differ with regard to parental investment and sexual selection strategies. Female strategies focus on obtaining the food necessary to sustain a pregnancy and choosing the best male(s) to father offspring. Male strategies focus on gaining access to receptive females.<\/p>\n<h3 class=\"import-Normal\"><strong>Parental Investment<\/strong><\/h3>\n<figure style=\"width: 362px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image9.jpg\" alt=\"Monkey holds baby.\" width=\"362\" height=\"241\" \/><figcaption class=\"wp-caption-text\">Figure 7.16: A female Japanese macaque nursing her infant. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Snow_monkey_baby_milk_time.jpg\">Snow monkey baby milk time<\/a> by Daisuke Tashiro 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>Biologically speaking, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1078\">parental investment<\/a><\/strong> is any time or energy a parent devotes to the current offspring that enhances its survival (and eventual reproductive success) at the expense of the parent\u2019s ability to invest in the next offspring (Trivers 1972). Female primates invest more heavily in offspring than males. Even before conception, females produce energy-containing eggs, and they will be responsible for sustaining a fertilized egg until it implants in the uterus. After that, they invest in pregnancy and lactation (Figure 7.16). Because all of this investment requires a lot of energy, female primates can only produce one offspring (or litter) at a time. A species\u2019 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1080\">interbirth interval<\/a><\/strong>, or the typical length of time between one birth and the next, is determined by the length of time necessary to maximize each offspring\u2019s survival without jeopardizing the female\u2019s ability to produce the greatest number of offspring possible. If a female invests too little (i.e., weans an offspring too early), she may give birth to many offspring, but very few (if any) of them will survive. If she invests too much (i.e., nurses an offspring even after it could be weaned), she ensures the survival of that individual offspring but will not be able to produce very many during her lifetime. To maximize her reproductive success, a female must invest <em>just<\/em> long enough to ensure the greatest number of offspring survive to reproduce. We often think of maternal care as an <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1082\">innate<\/a><\/strong> (or natural), instinctive behaviour. Yet this is not the case. The \u201cSpecial Topic: Is Maternal Behaviour Innate?\u201d dispels the myth that maternal behaviour is solely instinctual and explains how female primates learn to be good mothers.<\/p>\n<h3 class=\"import-Normal\"><strong>Sexual Selection<\/strong><\/h3>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1084\">Sexual selection<\/a><\/strong>, or selection for traits that maximize mating success, comes in two forms. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1086\">Intrasexual selection<\/a><\/strong> is selection for traits that enhance the ability of members of one sex to compete amongst themselves (\u201c<em>intra<\/em>sexual\u201d = within one sex). <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1088\">Intersexual selection<\/a><\/strong> is selection for traits that enhance the ability of one sex to attract the other (\u201c<em>inter<\/em>sexual\u201d = between the sexes).<\/p>\n<p class=\"import-Normal\">Intrasexual selection most often operates on males. In the wild, adult females are either pregnant or lactating for most of their adult lives. So, in a given population, there are usually more males available and willing to mate than there are females. The result? Females are a scarce resource over which males compete. Intrasexual selection favours traits that help a male win fights with other males. In primates, these traits include large body size (Figure 7.17a) and large canines (Figure 7.17b). Because females don\u2019t possess these same traits, males and females of some species look different; that is, they are sexually dimorphic (Figure 7.17a).<\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_207\" aria-describedby=\"caption-attachment-207\" style=\"width: 691px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-204\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.17.jpg\" alt=\"Male baboon with two females. Adult male baboon.\" width=\"691\" height=\"293\" \/><figcaption id=\"caption-attachment-207\" class=\"wp-caption-text\">Figure 7.17a-b: a. Hamadryas baboons (Papio hamadryas) are sexually dimorphic. The male (left) is much bigger than the female (centre) and also has different coloured fur. b. Adult males, like this gelada baboon, also have larger canines than females. Credit: a. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Dierenpark_Emmen_baboon_(2679944324).jpg\">Dierenpark Emmen baboon (2679944324)<\/a> by<a href=\"https:\/\/www.flickr.com\/people\/80538772@N00\"> robin bos<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\"> CC BY 2.0 License.<\/a> b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Olive_Baboon_Papio_anubis_in_Tanzania_3066_Nevit.jpg\">Olive Baboon Papio anubis, Picture Taken in Tanzania<\/a> by Nevit Dilmen has been modified (cropped) and 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\">Intersexual selection also tends to operate on males, selecting traits that make a male more attractive to females. Females, in turn, choose among potential fathers. Because female primates invest more in offspring production and care than males (see the \u201cParental Investment\u201d section, above), it is more costly for them if the offspring dies before maturity or reaches maturity but does not reproduce. Thus, it benefits a female primate to be choosy and try to pick the healthiest male as a mate. Males must display traits that tell a female why she should choose <em>him<\/em>, and not another male, as her mate.<\/p>\n<p class=\"import-Normal\">What traits are female primates looking for? In humans, women may look for a mate who can provide important resources, such as food, paternal care, or protection. This is rare in other primates, though, since most females do not need males to provide resources. More commonly, female primates obtain genetic benefits for their offspring from choosing one male over another. Often the specific criteria by which females select mates is unknown. However, if a female chooses a healthy (as indicated by traits like a plush coat, bright colouration, or large body size) or older male, she may obtain genes for her offspring that code for health or long life. If a male\u2019s rank is determined by competitive ability that has a genetic component, females who choose males who win fights may acquire these genes (and qualities) for their offspring. Females in some species appear to prefer new immigrants, sometimes even \u201csneaking\u201d copulations with males who are not established members of their groups. Such a preference may provide their offspring with novel genes and increase genetic variation (for more about the importance of genetic variation, see Chapter 4). Female choice is often more subtle than male-male competition, so it can be more difficult to study. However, as more research is conducted, we continue to improve our understanding of the ways that female primates exert their choice.<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Is Maternal Behaviour Innate?<\/h2>\n<p>Zoos almost always have nurseries where infants are cared for by zookeepers if their mothers will not care for them (Figure 7.18). These exhibits are among the most popular because the babies are so cute and so much fun to watch. And the caretaking positions in zoo nurseries are often among the most coveted by zoo personnel for the same reasons. But if maternal behaviour is instinctive, why do zoo nurseries even exist? The answer is that in many species, including primates, maternal behaviour is not purely instinctual; it is dependent on <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1090\">social learning<\/a><\/strong> (behaviour learned by observing and imitating others), as well.<\/p>\n<figure style=\"width: 433px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image13-1.png\" alt=\"Newborn orangutan feeding from a bottle.\" width=\"433\" height=\"336\" \/><figcaption class=\"wp-caption-text\">Figure 7.18: Newborn orangutan at Audubon Zoo being bottle-fed. Credit: <a href=\"https:\/\/newsroom.audubonnatureinstitute.org\/critically-endangered-orangutan-gives-birth-at-audubon-zoo\/\">Newborn orangutan born at Audubon Zoo being bottle fed<\/a> (2022) by<a href=\"https:\/\/audubonnatureinstitute.org\/\"> Audubon Nature Institute<\/a> is used by permission.<\/figcaption><\/figure>\n<p>Captive female primates, including gorillas and chimpanzees, who have not had the opportunity to observe their mother or other females care for infants do not know how to care for their own offspring. Although it is preferred that the primate mother care for her own infant, there are cases when she will not and humans must step in to ensure the offspring survives. When hand-rearing by humans is necessary, the infant is returned to the group as soon as possible in the hopes that it will learn species-typical behaviour from its mother and other conspecifics. Observations such as these indicate that maternal behaviour is learned, not innate, and that maternal care is critically important to the social and psychological development of young primates.<\/p>\n<\/div>\n<h2 class=\"import-Normal\">Communication<\/h2>\n<p class=\"import-Normal\">In its most basic form, communication occurs when one individual (the sender) emits a signal that conveys information, which is detected by another individual (the receiver). We have discussed several aspects of primate sociality in this chapter, all of which require the communication of information between individuals. But exactly <em>how<\/em> does a female chimpanzee communicate her sexual availability? <em>How<\/em> does a vervet monkey communicate the approach of a leopard or that a python is nearby? <em>How<\/em> do solitary, nocturnal primates, like the slow loris, communicate information about themselves to conspecifics? Primate communication comes in four forms: vocal, visual, olfactory, and tactile. Species vary in their reliance on each.<\/p>\n<h3 class=\"import-Normal\"><strong>Vocal Communication<\/strong><\/h3>\n<p class=\"import-Normal\">Primates use sound to communicate danger or threats, to claim and maintain a territory, or make contact with other group members. Alarm calls are given in response to predators. In some cases, alarm calls are used to alert members of the group to the presence of a predator so they can take evasive action. In other cases, they are directed at the predator itself, signaling that it has been detected. You can learn more about alarm calls as forms of vocal communication in the highlight box in this chapter entitled \u201cDig Deeper: Alarm Calls: Signals to Friends or Foes?.\u201d<\/p>\n<p class=\"import-Normal\">Loud calls are designed to travel great distances and are used in territorial defence by many primate species including indris (<em>Indri indri<\/em>), orangutans, gibbons, and howler monkeys (<em>Alouatta<\/em>). In dense forest, where visual communication can be difficult, loud calls can be useful in signaling to conspecifics that a group or individual occupies a specific area. Howler monkeys are named for their loud calls, or \u201croars,\u201d which can be heard one kilometre or more away (Sch\u00f6n Ybarra 1986). Howler monkey roars may act to maintain distance between neighbouring groups or keep extragroup males from entering the home range (1986).<\/p>\n<p class=\"import-Normal\">Other vocalizations are intended to communicate with individuals in one\u2019s own group. These include vocalizations given as part of threat displays or dominance interactions, as well as contact calls that provide information about one\u2019s location to other group members. Chacma baboons (<em>Papio ursinus<\/em>) have a rich repertoire of vocalizations for communicating with other group members (Fischer Et al. 2008). Adult males give specific vocalizations during threat displays and physical confrontations. Subordinates \u201cscreech\u201d when retreating from a dominant individual, signaling submission. Since baboons rely on membership in their group for finding food and detecting predators, a baboon separated from his group will vocalize in an attempt to regain contact. Young baboons emit their own contact calls when separated from their mothers.<\/p>\n<h3 class=\"import-Normal\"><strong>Visual Communication<\/strong><\/h3>\n<figure id=\"attachment_207-2\" aria-describedby=\"caption-attachment-207-2\" style=\"width: 493px\" class=\"wp-caption alignleft\"><img class=\"wp-image-206\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.19.jpg\" alt=\"Female baboon with sexual swelling. Male and female baboon.\" width=\"493\" height=\"209\" \/><figcaption id=\"caption-attachment-207-2\" class=\"wp-caption-text\">Figure 7.19a-b: Two female hamadryas baboons. The female on the left has a sexual swelling while the female on the right (in foreground, with infant clinging to her belly) does not. An adult male is behind her. Credit: a. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Sexual_swelling_in_female_Hamadryas_baboon.jpg\">Sexual swelling in female Hamadryas baboon<\/a> by<a href=\"https:\/\/www.flickr.com\/photos\/mamoritai\/\"> Mamoritai<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/legalcode\"> CC BY-SA 2.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Hamadryas_baboon_at_Giza_Zoo_by_Hatem_Moushir_36.JPG\">Hamadryas baboon at Giza Zoo by Hatem Moushir 36<\/a> by<a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Hatem_Moushir\"> Hatem Moushir<\/a> has been modified (cropped) and 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><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1092\">Visual communication<\/a><\/strong>, which involves signals that can be seen, is an important component of nonhuman primate behaviour, alone or in combination with other forms of communication. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1094\">Piloerection<\/a><\/strong>, or raising one\u2019s hair or fur, is used in aggressive interactions to make an individual appear larger than it actually is. Female macaques (<em>Macaca<\/em>), baboons (<em>Papio<\/em>), and chimpanzees, signal sexual receptivity through changes in the size, shape, and, often, colour of their hindquarters, called a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1096\">sexual swelling<\/a><\/strong> (Figure 7.19a). The sexual swelling reaches its maximum size at ovulation. When females are not receptive, either because they are pregnant or are nursing, they do not display a sexual swelling (Figure 7.19b). Thus, the presence or absence of a sexual swelling signals a female\u2019s reproductive state.<\/p>\n<p>Monkeys and apes use diverse facial expressions in visual communication. Showing your teeth in a \u201csmile\u201d sends a signal of friendship in humans. Displaying teeth in this way is a sign of anxiety or fear in primates. That male mandrill you see \u201cyawning\u201d at your local zoo is actually displaying his teeth to signal tension or to threaten a rival (Figure 7.20a). In addition to showing their canines, male gelada baboons use \u201clip flips,\u201d in which the gums and teeth are exposed by flipping the upper lip up over the nostrils (Figure 7.20b), and \u201craised eyelids,\u201d in which the pale eyelids are exposed by pulling the scalp back as threatening gestures (Aich, Moos-Heilen, &amp; Zimmerman 1990). Submissive males respond by fleeing or presenting their hindquarters.<\/p>\n<figure id=\"attachment_207-3\" aria-describedby=\"caption-attachment-207-3\" style=\"width: 597px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-207\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.20-1.jpg\" alt=\"Adult male mandrill and adult male hamadryas baboon yawning.\" width=\"597\" height=\"310\" \/><figcaption id=\"caption-attachment-207-3\" class=\"wp-caption-text\">Figure 7.20a-b: Males use visual displays to communicate with other males. The male mandrill (left) is yawning to display his canines, and the male gelada baboon (right) enhances the yawn by flipping his upper lip back and raising his eyelids. Credit: a. <a href=\"https:\/\/pxhere.com\/en\/photo\/559944\">Mandrill<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/mathiasappel\/with\/24639723420\/\">Mathias Appel<\/a> has been modified (cropped) and designated to the <a href=\"https:\/\/creativecommons.org\/share-your-work\/public-domain\/cc0\/\">public domain (CC0)<\/a>. b.<a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:BabouinGeladaAuReveil.JPG\"> BabouinGeladaAuReveil<\/a> by<a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:BluesyPete\"> BluesyPete<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\"> CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<figure id=\"attachment_209\" aria-describedby=\"caption-attachment-209\" style=\"width: 414px\" class=\"wp-caption alignleft\"><img class=\"wp-image-208\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.22.jpg\" alt=\"A bald uakari. A spider monkey.\" width=\"414\" height=\"154\" \/><figcaption id=\"caption-attachment-209\" class=\"wp-caption-text\">Figure 7.22a-b: Many monkey species have colourful faces, including the bald uakari (Cacajao calvus; left) and the white-bellied spider monkey (Ateles belzebuth) (right). Credit: a. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Uakari.jpg\">Uakari<\/a> by Coada dragos has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\"> CC BY-SA 4.0 License<\/a>. 6.22b <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Ateles_belzebuth_(White-bellied_spider_monkey)_2.jpg\">Ateles belzebuth (White-bellied spider monkey) 2<\/a> by<a href=\"https:\/\/www.flickr.com\/photos\/ewas-world\/\"> Ewa<\/a> (username: Ewcek65) has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\"> CC BY 2.0 License<\/a>.<\/figcaption><\/figure>\n<figure style=\"width: 163px\" class=\"wp-caption alignright\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-3.png\" alt=\"A male mandrill\u2019s face.\" width=\"163\" height=\"246\" \/><figcaption class=\"wp-caption-text\">Figure 7.21: The colourful face of the male mandrill provides information about health and fitness to other mandrills.Credit: <a href=\"https:\/\/www.flickr.com\/photos\/mathiasappel\/24842082402\/\">Mandrill<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/mathiasappel\/with\/24639723420\/\">Mathias Appel<\/a> has been modified (cropped) and designated to the <a href=\"https:\/\/creativecommons.org\/share-your-work\/public-domain\/cc0\/\">public domain (CC0)<\/a>.<\/figcaption><\/figure>\n<p>Primates also communicate through colour. In female and male mandrills, facial colouration provides information about an individual\u2019s health, competitive ability, and reproductive state to conspecifics (Figure 7.21; Setchell &amp;t al. 2008; Setchell, Wickings, &amp; Knapp 2006). Variation in facial colouration among monkeys of Central and South America ranges from very simple (Figure 7.22a) to complex (Figure 7.22b). Species living with larger numbers of other primate species have evolved more complex facial colouration patterns, suggesting that this trait evolved as a form of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1098\">species recognition<\/a><\/strong>, or the ability to differentiate conspecifics from members of other species (Santana, Lynch Alfaro, &amp; Alfaro 2012).<\/p>\n<h3 class=\"import-Normal\"><strong>Olfactory Communication<\/strong><\/h3>\n<p class=\"import-Normal\">All primates use scent to communicate. Females secrete chemicals from their <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1100\">anogenital<\/a> <\/strong>region (the area of the anus and genitals) that provide males with information about their reproductive state. In some species, like macaques and chimpanzees, this olfactory signal is enhanced by a sexual swelling, as discussed above. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1102\">Olfactory communication<\/a><\/strong>, or communicating through scent, is particularly important for monkeys of Central and South America, lemurs, and lorises. Male and female common squirrel monkeys (<em>Saimiri sciureus<\/em>) (Figure 7.23a) engage in \u201curine washing,\u201d in which an individual urinates on its hands and feet and then uses them to spread urine all over its body. Urine washing may be used to mark trails for others to follow, to control body temperature, as part of dominance displays, or to communicate reproductive state (Boinski 1992). During aggressive interactions with other males, male ring-tailed lemurs rub their tails with scent from glands on their wrists and chests. They use their \u201cperfumed\u201d tails in aggressive interactions with other males, who may respond by waiving their own scented tail, with physical aggression, or by fleeing (Jolly 1966). Males also waive their tails, saturated in scent, to attract females (Shirasu et al. 2020). Males use scent glands in their wrists to mark territorial boundaries (Figure 7.23b; Mertl-Millhollen 1988).<\/p>\n<\/div>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_210-7\" aria-describedby=\"caption-attachment-210-7\" style=\"width: 641px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-210\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.23.jpg\" alt=\"A squirrel monkey. A ring-tailed lemur.\" width=\"641\" height=\"332\" \/><figcaption id=\"caption-attachment-210-7\" class=\"wp-caption-text\">Figure 7.23a-b: Some primates, like the common squirrel monkey (left) and the ring-tailed lemur (right), communicate using scent. Credit: a. <a href=\"https:\/\/www.flickr.com\/photos\/rubund\/6337874822\/\">Saimiri sciureus<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/rubund\/\">Ruben Undheim<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/\">CC BY-SA 2.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Lemur_catta_004.jpg\">Lemur catta 004<\/a> by <a href=\"https:\/\/en.wikipedia.org\/wiki\/User:Maky\">Maky<\/a> has been modified (cropped) and 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<div class=\"__UNKNOWN__\">\n<h3 class=\"import-Normal\"><strong>Tactile Communication<\/strong><\/h3>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1104\">Tactile communication<\/a><\/strong>, or communicating through touch, is very important in all primate species. Physical contact is used to comfort and reassure, is part of courtship and mating, and is used to establish dominance and alliances. Grooming is an important and clearly enjoyable form of tactile communication for all primates (Figure 7.24). Not only does grooming serve to clean the skin and fur, removing parasites and debris, but it is an important affiliative behaviour that helps reinforce social bonds, repair relationships, and cement alliances.<\/p>\n<figure style=\"width: 674px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image1-8.png\" alt=\"Four primate species grooming.\" width=\"674\" height=\"478\" \/><figcaption class=\"wp-caption-text\">Figure 7.24: Examples of grooming in Japanese macaques (upper left), tufted capuchins (Sapajus apella) (upper right), gelada baboons (lower left), and black-and-white ruffed lemurs (Varecia variegata; lower right). Credit: Examples of grooming original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Jaffe 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:Yakushima_macaques_grooming_each_other.jpg\">Yakushima macaques grooming each other<\/a> by<a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Grendelkhan\"> Grendelkhan<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Tufted_capuchin_monkeys_grooming_session_III.jpg\">Tufted capuchin monkeys grooming session III<\/a> by Adrian Soldati, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>;<a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Baboons_Wunania_012018.jpg\"> Baboons Wunania 012018<\/a> by Kim Toogood, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>;<a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Black-and-white_ruffed_lemur_03.jpg\"> Black-and-white ruffed lemur 03<\/a> by Mattis2412, <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/deed.en\">public domain (CC0 1.0)<\/a>].<\/figcaption><\/figure>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Dig Deeper: Alarm Calls: Signals to Friends or Foes?<\/h2>\n<p class=\"import-Normal\">Alarm calls are common among group-living primates. They often serve to notify conspecifics of potential danger, as is the case with vervet monkeys. Research has shown that: (1) vervets classify predators based on hunting style; (2) alarm calls convey information to other vervets about that hunting style; and (3) other vervets respond in ways appropriate for evading that type of predator (Seyfarth, Cheney, &amp; Marler 1980a). When a vervet gives a \u201cleopard\u201d alarm call (directed at mammalian carnivores like leopards, Figure 7.25a), monkeys on the ground climb the nearest tree, while monkeys already in trees stay there or climb higher. Since most mammalian carnivores hunt on the ground, getting into, and staying in, a tree is the best option for escape. When the \u201csnake\u201d alarm call is given, vervets stand on their hind legs and look down at the ground (Figure 7.25b). Since snakes are not pursuit predators, locating them quickly so as to avoid them is the best strategy. Lastly, when an \u201ceagle\u201d alarm call is given, vervets look up or run into bushes, both of which are useful responses for avoiding hawks and eagles, which attack from above (Figure 7.25c). Vervets clearly understand the meaning of each type of alarm call, as they respond appropriately even when they do not see the actual predator (Seyfarth, Cheney, &amp; Marler 1980b). Such <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1106\">semantic communication<\/a><\/strong>, which involves the systematic use of signals to refer to objects in the environment, was once believed to be unique to humans. It may be a precursor to the symbolic capacities of human language.<\/p>\n<figure style=\"width: 482px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5-1.jpg\" alt=\"Primate in a tree views a leopard.\" width=\"482\" height=\"344\" \/><figcaption class=\"wp-caption-text\">Figure 7.25a<\/figcaption><\/figure>\n<figure style=\"width: 484px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image2-2.jpg\" alt=\"Primate views snake on the ground.\" width=\"484\" height=\"338\" \/><figcaption class=\"wp-caption-text\">Figure 7.25b<\/figcaption><\/figure>\n<figure style=\"width: 481px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-2.jpg\" alt=\"Primate on the ground sees bird.\" width=\"481\" height=\"517\" \/><figcaption class=\"wp-caption-text\">Figure 7.25c\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 Figure 7.25a-c: Vervet monkeys respond in different ways to alarm calls for each of their three main predators (leopards, snakes, and eagles) which are appropriate to predator hunting strategies. Credit: Vervet Monkey Alarm Calls by Mary Nelson, original to Explorations: An Open Invitation to Biological Anthropology, 2nd edition, 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\">Research on other African monkeys indicates that some species use alarm calls to signal to the predator that it has been detected. Diana monkeys (<em>Cercopithecus diana<\/em>) give alarm calls to leopards (<em>Panthera pardus<\/em>) but not chimpanzees (Zuberb\u00fchler, No\u00eb, &amp; Seyfarth 1997). Because leopards are stealth predators, they rely on the element of surprise to sneak up on their prey (Figure 7.26a). Alarm calling at leopards appears to tell the leopard that it has been seen and therefore its chance of success will be low. Research shows leopards are more likely to stop hunting after an alarm call has been emitted. Unlike leopards, chimpanzees are pursuit predators and may even use alarm calls to locate potential prey (Figure 7.26b). With such a predator, prey are better off remaining as silent as possible so as not to alert the predator to their location (Zuberb\u00fchler Et al. 1999).<\/p>\n<figure id=\"attachment_215\" aria-describedby=\"caption-attachment-215\" style=\"width: 1907px\" class=\"wp-caption alignnone\"><img class=\"wp-image-215 size-full\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.26.jpg\" alt=\"Leopard crouches in grass. Chimpanzee looks up. \" width=\"1907\" height=\"589\" \/><figcaption id=\"caption-attachment-215\" class=\"wp-caption-text\">Figure 7.26a-b: Because leopards (left) and chimpanzees (right) hunt differently, Diana monkeys react differently to them. Credit: a. <a href=\"https:\/\/www.flickr.com\/photos\/thimindu\/5842997328\">Crouching Leopard<\/a> by<a href=\"https:\/\/www.flickr.com\/photos\/thimindu\/\"> Thimindu Goonatillake<\/a> is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/\"> CC BY-SA 2.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Chimpanzee_in_the_wild.jpg\">Chimpanzee in the wild<\/a> by D.G. Kulakov 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\">\n<\/div>\n<h2 class=\"import-Normal\"><span style=\"text-align: initial;font-size: 1em\">The Question of Culture<\/span><\/h2>\n<p class=\"import-Normal\">It may be surprising in a chapter on nonhuman primates to see a discussion of culture. After all, culture is considered by many, including cultural anthropologists, to be a distinguishing characteristic of humans. Indeed, some anthropologists question claims of culture in primates and other animals. Definitions of animal culture focus on specific behaviours that are unique to one population. Anthropological definitions of human culture emphasize shared ideology (e.g., values, morals, beliefs) and symbols, not just behaviour. Using this definition, some cultural anthropologists view primates as lacking culture because of the absence of symbolic life (e.g., religion). However, the longer we study primate groups and populations, the more insight we gain into primate behavioural variation. If we define <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1108\">culture<\/a><\/strong> as the transmission of behaviour from one generation to the next through social learning, then we must view at least some of the behavioural variation we see in primates as forms of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1110\">cultural tradition<\/a><\/strong>, or a distinctive pattern of behaviour shared by multiple individuals in a social group that persists over time (Whiten 2001).<\/p>\n<h3 class=\"import-Normal\"><strong>Chimpanzee Culture<\/strong><\/h3>\n<p class=\"import-Normal\">Due to both their high level of intelligence and the large number of long-term studies on several different populations, chimpanzees provide the best example of cultural tradition in primates. Chimpanzees express cultural variation in multiple behavioural patterns, ranging from population-specific prey preferences and hunting strategies to tool-use techniques and social behaviours. For example, in Tanzania, chimpanzees fish for termites by stripping twigs and then poking the twigs into termite mounds. The termites react to the \u201cinvasion\u201d by attacking the twig. The chimpanzee pulls the twig out, termites attached, and eats them. In Gambia, they use modified twigs to extract honey from holes in trees. In Fongoli, S\u00e9n\u00e9gal, chimpanzees use sticks as \u201cspears\u201d that they stab into tree cavities to hunt for galagos (Figure 7.27). Multiple chimpanzee populations use a \u201chammer and anvil\u201d to crack open nuts, but the specific techniques differ. Because the cultural traditions are so diverse and unique, if a researcher can observe enough of a chimpanzee\u2019s behaviour, it is possible to assign that individual to a specific community, much in the same way a human being can be associated with a specific culture based on his or her behaviour (Whiten 2011).<\/p>\n<figure style=\"width: 800px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image3-4.jpg\" alt=\"Chimpanzees hunting galagos by poking them with a stick. \" width=\"800\" height=\"453\" \/><figcaption class=\"wp-caption-text\">Figure 7.27a-d: Tool-assisted hunting by a chimpanzee at Fongoli, S\u00e9n\u00e9gal. An adult male chimpanzee uses a tree branch with a modified end to (a\u2013c) stab into a cavity within a hollow tree branch that houses a galago. He ultimately captures the galago as (d) his adolescent brother looks on. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Pan_troglodytes,_tool_use_in_Senegal.jpg\">Pan troglodytes, tool use in Senegal<\/a> by<a href=\"https:\/\/royalsocietypublishing.org\/content\/2\/4\/140507\"> J. D. Pruetz, P. Bertolani, K. Boyer Ontl, S. Lindshield, M. Shelley, and E. G. Wessling<\/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\">How do chimpanzee cultures develop, and how does cultural transmission occur? Although we do not know for sure how chimpanzee cultural traditions develop initially, it is possible that different groups invent, either accidentally or deliberately, certain behaviours that other individuals copy. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1112\">Immigration<\/a><\/strong>, or movement of an individual into a new group or community, is an important avenue of cultural transmission in chimpanzees, much as it is between human cultures. Immigrants (typically females) may bring cultural traditions to their new community, which residents observe and learn. Conversely, immigrants may observe and learn a cultural tradition practiced in their new community (Whiten 2011).<\/p>\n<h3 class=\"import-Normal\"><strong>Cultural Transmission in Macaques<\/strong><\/h3>\n<p class=\"import-Normal\">Two monkey species are well-known for behavioural variation that has been called \u201cpre-cultural\u201d by some primatologists: Japanese macaques and tufted capuchins (<em>Sapajus apella<\/em>). The transmission of unique <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1114\">foraging<\/a><\/strong> (the act of searching for food) behaviours through the members of a provisioned group of Japanese macaques on Koshima Island is well known (Matsuzawa 2015). In an effort to keep the monkeys nearby, researchers provided them with piles of sweet potatoes. A juvenile female named Imo spontaneously washed a muddy sweet potato in a stream. This new food-processing technique first spread among other juveniles and then gradually to older individuals. Within 30 years, it had spread across generations, and 46 of 57 monkeys in the group engaged in the behaviour. Another example comes from a group living far to the north, in Shiga-Heights, Nagano Prefecture. Researchers used apples to entice Japanese macaques to the area. Within a few years, monkeys visited the area regularly and were observed playing with the water in the hot springs. Soon, they climbed into the hot springs and learned to immerse themselves to keep warm and reduce stress when not foraging (Figure 7.28; Matsuzawa 2018; Takeshita Et al. 2018; recall also our discussion of hot spring use as an example of analogous traits at the beginning of this chapter). These examples share several characteristics with human culture, including invention or modification of behaviour, transmission of behaviour between individuals, and the persistence of the behaviour across generations (McGrew 1998).<\/p>\n<figure style=\"width: 477px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image6-4.png\" alt=\"Two monkeys in a hot spring.\" width=\"477\" height=\"292\" \/><figcaption class=\"wp-caption-text\">Figure 7.28: Hot spring use by Japanese macaques is a culturally transmitted behaviour. Credit: <a href=\"https:\/\/www.flickr.com\/photos\/peterthoeny\/32160301021\">Oooh, This Feels Sooo Good!<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/peterthoeny\/\">Peter Theony - Quality HD Photography<\/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<h2 class=\"import-Normal\">Summary<\/h2>\n<p class=\"import-Normal\">Primates are socially complex, extremely intelligent, and highly adaptable. In this chapter we discussed aspects of primate ecology, including how body size and characteristics of food affect what primates eat and how primates interact with other species in their environment. We examined why primates live in groups, the types of groups in which they are found, and the reproductive strategies used by males and females to maximize reproductive success. Like other aspects of their behaviour, primate communication is varied and complex, and we discussed how primates communicate using vocal, visual, olfactory, and tactile signals. Finally, we explored the question of culture among nonhuman primates and learned that some species have cultural traditions, distinctive patterns of behaviour shared by multiple individuals in a social group that persist over time. Humans and other primates are similar in many ways. Learning about principles of primate ecology and behaviour can help us better understand our own behaviour and the behaviours of our extinct relatives.<\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Review Questions<\/h2>\n<ul>\n<li class=\"import-Normal\">If anthropology is the study of humans, why do some anthropologists study primates?<\/li>\n<li class=\"import-Normal\">How does a primate\u2019s ecology affect their diet and interactions with other organisms?<\/li>\n<li class=\"import-Normal\">Why do primates live in groups and in what types of groups do they live?<\/li>\n<li class=\"import-Normal\">What is parental investment and sexual selection?<\/li>\n<li class=\"import-Normal\">What are some examples of primate communication?<\/li>\n<li class=\"import-Normal\">What is the evidence for cultural traditions in primates and how do primatologists think cultural transmission occurs in primates?<\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\">Key Terms<\/h2>\n<p class=\"import-Normal\"><strong>Abundance<\/strong>: How much food is available in a given area.<\/p>\n<p class=\"import-Normal\"><strong>Adaptation<\/strong>: A trait with a function.<\/p>\n<p class=\"import-Normal\"><strong>Affiliative<\/strong>: Nonaggressive social interactions and associations between individuals.<\/p>\n<p class=\"import-Normal\"><strong>Agonistic<\/strong>: Conflict; aggressive interactions between individuals.<\/p>\n<p class=\"import-Normal\"><strong>Alarm calling<\/strong>: Vocalizations emitted by social animals in response to danger.<\/p>\n<p class=\"import-Normal\"><strong>Analogy<\/strong>: A similar trait found in different species that arose independently.<\/p>\n<p class=\"import-Normal\"><strong>Anogenital<\/strong>: Relating to the anus and genitals.<\/p>\n<p class=\"import-Normal\"><strong>Breeding season<\/strong>: The time of year when females are receptive to mating.<\/p>\n<p class=\"import-Normal\"><strong>Callitrichids<\/strong>: The primate family that includes marmosets and tamarins.<\/p>\n<p class=\"import-Normal\"><strong>Carnivores<\/strong>: Organisms whose diet consists primarily of animal tissue.<\/p>\n<p class=\"import-Normal\"><strong>Coalition<\/strong>: A temporary alliance between individuals.<\/p>\n<p class=\"import-Normal\"><strong>Community ecology<\/strong>: The branch of ecology that deals with the relationships and interactions between different organisms that occupy the same habitat.<\/p>\n<p class=\"import-Normal\"><strong>Comparison<\/strong>: An examination of the similarities and differences between two things, such as two primate species.<\/p>\n<p class=\"import-Normal\"><strong>Conspecifics<\/strong>: Members of the same species.<\/p>\n<p class=\"import-Normal\"><strong>Cooperative breeding<\/strong>: When individuals other than the mother and father help raise the offspring.<\/p>\n<p class=\"import-Normal\"><strong>Crypsis<\/strong>: The ability to avoid detection by other organisms, such as predators.<\/p>\n<p class=\"import-Normal\"><strong>Cultural tradition<\/strong>: A distinctive pattern of behaviour shared by multiple individuals in a social group, which persists over time and is acquired through social learning.<\/p>\n<p class=\"import-Normal\"><strong>Culture<\/strong>: The transmission of behaviour from one generation to the next through observation and imitation.<\/p>\n<p class=\"import-Normal\"><strong>Decolonize<\/strong>: Understanding and highlighting the theory and research of non-Western individuals and perspectives.<\/p>\n<p class=\"import-Normal\"><strong>Descendant<\/strong>: A species that comes after the ancestor species.<\/p>\n<p class=\"import-Normal\"><strong>Direct competition:<\/strong> Competition that involves physical interaction between individuals, such as fighting.<\/p>\n<p class=\"import-Normal\"><strong>Dispersal<\/strong>: To leave one\u2019s group or area. This may or may not involve joining another group.<\/p>\n<p class=\"import-Normal\"><strong>Distribution<\/strong>: How food is spread out.<\/p>\n<p class=\"import-Normal\"><strong>Diurnal<\/strong>: Active during the day.<\/p>\n<p class=\"import-Normal\"><strong>Dominance hierarchy<\/strong>: The ranked organization of individuals established by the outcome of aggressive-submissive interactions.<\/p>\n<p class=\"import-Normal\"><strong>Dominant<\/strong>: Being of high rank.<\/p>\n<p class=\"import-Normal\"><strong>Ecology<\/strong>: The relationship between organisms and their physical surroundings.<\/p>\n<p class=\"import-Normal\"><strong>Ecotourism<\/strong>: A form of tourism that focuses on nature-based attractions to provide learning opportunities and that uses economically and ecologically sustainable practices.<\/p>\n<p class=\"import-Normal\"><strong>Ethology<\/strong>: The study of animal behaviour.<\/p>\n<p class=\"import-Normal\"><strong>Fission-fusion<\/strong>: Societies in which group composition is flexible, such as chimpanzee and spider monkey societies. Individuals may break up into smaller feeding groups (fission) and combine into larger groups (fusion).<\/p>\n<p class=\"import-Normal\"><strong>Fitness<\/strong>: An individual\u2019s ability to survive and reproduce relative to other members of the same species.<\/p>\n<p class=\"import-Normal\"><strong>Folivores<\/strong>: Organisms whose diet consists primarily of leaves.<\/p>\n<p class=\"import-Normal\"><strong>Foraging<\/strong>: The act of searching for food.<\/p>\n<p class=\"import-Normal\"><strong>Frugivores<\/strong>: Organisms whose diet consists primarily of fruit.<\/p>\n<p class=\"import-Normal\"><strong>Grooming<\/strong>: Picking through the fur of another individual for cleaning or bonding purposes.<\/p>\n<p class=\"import-Normal\"><strong>Heterospecifics<\/strong>: Members of different species.<\/p>\n<p class=\"import-Normal\"><strong>Holism<\/strong>: The idea that the parts of a system interconnect and interact to make up the whole.<\/p>\n<p class=\"import-Normal\"><strong>Home range<\/strong>: The area that a group or individual uses over a given period of time (often over a year).<\/p>\n<p class=\"import-Normal\"><strong>Homology<\/strong>: A similar trait found in different species because it was inherited from a common ancestor.<\/p>\n<p class=\"import-Normal\"><strong>Immigration<\/strong>: Movement of an individual into a new group or community.<\/p>\n<p class=\"import-Normal\"><strong>Indirect competition<\/strong>: Competition that does not involve physical interaction between individuals, such as eating food before another individual arrives at the food site.<\/p>\n<p class=\"import-Normal\"><strong>Infanticide<\/strong>: The killing of infants of one\u2019s own species.<\/p>\n<p class=\"import-Normal\"><strong>Innate<\/strong>: Natural; as in behaviour that comes naturally.<\/p>\n<p class=\"import-Normal\"><strong>Insectivores<\/strong>: Organisms whose diets consist primarily of insects.<\/p>\n<p class=\"import-Normal\"><strong>Interbirth interval<\/strong>: The typical length of time between one birth and the next for a species.<\/p>\n<p class=\"import-Normal\"><strong>Intersexual selection<\/strong>: The selection for traits that enhance the ability of the members of one sex to attract the attention of the other.<\/p>\n<p class=\"import-Normal\"><strong>Intrasexual selection<\/strong>: Selection for traits that enhance the ability of members of one sex to compete amongst themselves.<\/p>\n<p class=\"import-Normal\"><strong>Mating system<\/strong>: A way of describing which male(s) and female(s) mate.<\/p>\n<p class=\"import-Normal\"><strong>Metabolism<\/strong>: The chemical changes that take place in an organism that turn nutrients into energy.<\/p>\n<p class=\"import-Normal\"><strong>Mobbing<\/strong>: Cooperatively attacking or harassing a predator.<\/p>\n<p class=\"import-Normal\"><strong>Monogamy<\/strong>: A mating system in which one male mates with one female.<\/p>\n<p class=\"import-Normal\"><strong>Multi-male, multi-female<\/strong>: A group that consists of multiple adult males, multiple adult females, and their dependent offspring.<\/p>\n<p class=\"import-Normal\"><strong>Multi-male, single-female<\/strong>: A group that consists of two or more adult males, one breeding female, their dependent offspring, and non-breeding females.<\/p>\n<p class=\"import-Normal\"><strong>Mutualistic\/mutualism<\/strong>: When different species work together, with each benefiting from the interaction.<\/p>\n<p class=\"import-Normal\"><strong>Niche<\/strong>: The role of a species in its environment; how it meets its needs for food, shelter, etc.<\/p>\n<p class=\"import-Normal\"><strong>Nocturnal<\/strong>: Active at night.<\/p>\n<p class=\"import-Normal\"><strong>Olfactory communication<\/strong>: Conveying information through scent.<\/p>\n<p class=\"import-Normal\"><strong>Omnivores<\/strong>: Organisms whose diet consists of plant and animal matter.<\/p>\n<p class=\"import-Normal\"><strong>Pair bond<\/strong>: A strong, long-term relationship between two individuals.<\/p>\n<p class=\"import-Normal\"><strong>Parasite<\/strong>: An organism that lives in or on another organism.<\/p>\n<p class=\"import-Normal\"><strong>Parental investment<\/strong>: Any time or energy a parent devotes to the current offspring that enhances its survival (and eventual reproductive success) at the expense of the parent\u2019s ability to invest in the next offspring.<\/p>\n<p class=\"import-Normal\"><strong>Philopatric<\/strong>: Remaining in the group of one\u2019s birth.<\/p>\n<p class=\"import-Normal\"><strong>Piloerection<\/strong>: Raising one\u2019s hair or fur in an effort to look bigger.<\/p>\n<p class=\"import-Normal\"><strong>Polyandry<\/strong>: A mating system in which multiple males mate with a single breeding female.<\/p>\n<p class=\"import-Normal\"><strong>Polygamy<\/strong>: A mating system in which multiple males mate with multiple females.<\/p>\n<p class=\"import-Normal\"><strong>Polygyny<\/strong>: A mating system in which one male mates with multiple females.<\/p>\n<p class=\"import-Normal\"><strong>Polyspecific association<\/strong>: An association between two or more different species that involves behavioural changes in at least one of them to maintain the association.<\/p>\n<p class=\"import-Normal\"><strong>Primate community<\/strong>: All primate species that occur in an area.<\/p>\n<p class=\"import-Normal\"><strong>Primatologist<\/strong>: A scientist who studies primate behaviour and\/or ecology.<\/p>\n<p class=\"import-Normal\"><strong>Primatology<\/strong>: The scientific field that studies primate behaviour and\/or ecology.<\/p>\n<p class=\"import-Normal\"><strong>Ranging behaviour<\/strong>: Refers to the way in which animals move about their environment.<\/p>\n<p class=\"import-Normal\"><strong>Receptive<\/strong>: A term used to describe females who are ready for sexual reproduction (i.e., not pregnant or nursing).<\/p>\n<p class=\"import-Normal\"><strong>Reproductive success<\/strong>: An individual\u2019s genetic contribution to future generations, often measured through the number of offspring produced.<\/p>\n<p class=\"import-Normal\"><strong>Reproductive suppression<\/strong>: The prevention or inhibition of reproduction of healthy adults.<\/p>\n<p class=\"import-Normal\"><strong>Resident male<\/strong>: Term that describes the male who lives with a group of females.<\/p>\n<p class=\"import-Normal\"><strong>Seed dispersal<\/strong>: The process by which seeds move away from the plant that produced them in preparation for germination and becoming a new plant.<\/p>\n<p class=\"import-Normal\"><strong>Semantic communication<\/strong>: The systematic use of signals to refer to objects in the environment.<\/p>\n<p class=\"import-Normal\"><strong>Sexual dimorphism<\/strong>: When males and females of a species have different morphological traits.<\/p>\n<p class=\"import-Normal\"><strong>Sexual selection<\/strong>: The selection for traits that increase mating success. This occurs via intersexual selection and intrasexual selection.<\/p>\n<p class=\"import-Normal\"><strong>Sexual swelling<\/strong>: Area of the hindquarters that change in size, shape, and often colour over the course of a female\u2019s reproductive cycle, reaching maximum size at ovulation. Occurs in many primate species that live in Africa and Asia.<\/p>\n<p class=\"import-Normal\"><strong>Sexually monomorphic<\/strong>: When males and females of a species have similar morphological traits.<\/p>\n<p class=\"import-Normal\"><strong>Single-male, multi-female<\/strong>: A group that consists of one adult male, multiple adult female, and their dependent offspring.<\/p>\n<p class=\"import-Normal\"><strong>Single-male, single-female<\/strong>: A group that consists of one adult male, one adult female, and their dependent offspring.<\/p>\n<p class=\"import-Normal\"><strong>Social learning<\/strong>: The idea that new behaviours can be acquired by observing and imitating others.<\/p>\n<p class=\"import-Normal\"><strong>Social system<\/strong>: A way of describing the typical number of males and females of all age classes that live together.<\/p>\n<p class=\"import-Normal\"><strong>Social transmission<\/strong>: Transfer of something from one individual to another; this can include parasites, information, or cultural traditions.<\/p>\n<p class=\"import-Normal\"><strong>Sociality<\/strong>: The tendency to form social groups.<\/p>\n<p class=\"import-Normal\"><strong>Solitary<\/strong>: Living alone.<\/p>\n<p class=\"import-Normal\"><strong>Species recognition<\/strong>: The ability to differentiate conspecifics from members of other species.<\/p>\n<p class=\"import-Normal\"><strong>Subordinate<\/strong>: Being of low rank.<\/p>\n<p class=\"import-Normal\"><strong>Tactile communication<\/strong>: Conveying information through touch.<\/p>\n<p class=\"import-Normal\"><strong>Territory:<\/strong> A home range whose boundary is defended from intrusion by conspecifics.<\/p>\n<p class=\"import-Normal\"><strong>Vertebrates<\/strong>: The group of animals characterized by an internal spinal column or backbone. This includes fish, amphibians, reptiles, birds, and mammals.<\/p>\n<p class=\"import-Normal\"><strong>Vigilance<\/strong>: Watchful behaviour used to detect potential danger, usually in the form of predators or potential competitors.<\/p>\n<p class=\"import-Normal\"><strong>Visual communication<\/strong>: Conveying information through signals that can be seen.<\/p>\n<p class=\"import-Normal\"><strong>Vocal communication<\/strong>: Conveying information through signals that can be heard.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration<\/h2>\n<p class=\"import-Normal\">Goodall, Jane. 1971. <em>In the Shadow of Man<\/em>. Boston: Houghton Mifflin.<\/p>\n<p class=\"import-Normal\">Rowe, Noel, and Marc Myers, eds. 2016. <em>All the World\u2019s Primates. <\/em>Charleston, RI: Pogonias Press.<\/p>\n<p class=\"import-Normal\">Strier, Karen B. 2017. <em>Primate Behavioural Ecology.<\/em> 5th ed. New York: Routledge.<\/p>\n<p class=\"import-Normal\"><a href=\"https:\/\/pin.primate.wisc.edu\/\">Primate Info Net<\/a> is an information service of the National Primate Research Center at the University of Wisconsin, Madison. It includes Primate Factsheets, primate news and publications, a list of primate-related jobs, and an international directory of primatology, among other information.<\/p>\n<p class=\"import-Normal\"><a href=\"https:\/\/www.primate-sg.org\/\">Primate Specialist Group<\/a> is a collection of scientists and conservationists who work in dozens of African, Asian, and Latin American nations to promote research on primate conservation.<\/p>\n<p class=\"import-Normal\">Short videos of some primate behaviours discussed in this chapter:<\/p>\n<ul>\n<li class=\"import-Normal\">Watch vervet monkeys respond to different types of predators: BBC One. n.d. \u201cVervet Monkey\u2019s Escape Plans - Talk to the Animals: Episode 2 Preview.\u201d Accessed December 16, 2022. <a class=\"rId10\" href=\"https:\/\/www.youtube.com\/watch?v=q8ZG8Dpc8mM\">https:\/\/www.youtube.com\/watch?v=q8ZG8Dpc8mM. <\/a><\/li>\n<li class=\"import-Normal\">Watch male gelada baboons use the lip flip in competition with other males: Smithsonian Channel, June 9, 2017. \u201cWhy These Vegetarian Monkeys Have Sharp Predator Teeth.\u201d Accessed July 25, 2019. <a class=\"rId11\" href=\"https:\/\/www.youtube.com\/watch?time_continue=145&amp;v=aC6iYj_EBjY\">https:\/\/www.youtube.com\/watch?time_continue=145&amp;v=aC6iYj_EBjY<\/a>.<\/li>\n<li class=\"import-Normal\">Watch (and listen to!) howler monkeys \u201croar\u201d: Science News. N.d. \u201cHear a Male Howler Monkey Roar.\u201d Accessed November 21, 2022. <a class=\"rId12\" href=\"https:\/\/www.youtube.com\/watch?v=PYar0dkZ6v8\">https:\/\/www.youtube.com\/watch?v=PYar0dkZ6v8<\/a>.<\/li>\n<li class=\"import-Normal\">Watch Japanese macaques using natural hot springs: National Geographic. N.d. \u201cMeditative Snow Monkeys Hang Out in Hot Springs.\u201d Accessed July 25, 2019. <a class=\"rId13\" href=\"https:\/\/www.youtube.com\/watch?v=Aat9O85ynsI\">https:\/\/www.youtube.com\/watch?v=Aat9O85ynsI<\/a>.<\/li>\n<li class=\"import-Normal\">Watch chimpanzees make and use tools: National Geographic. n.d. \u201cChimps and Tools.\u201d Accessed July 25, 2019. <a class=\"rId14\" href=\"https:\/\/www.youtube.com\/watch?v=o2TBicMRLtA\">https:\/\/www.youtube.com\/watch?v=o2TBicMRLtA<\/a>.<\/li>\n<\/ul>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\">Aich, H., R. Moos-Heilen, and E. Zimmermann. 1990. \u201cVocalizations of Adult Gelada Baboons (<em>Theropithecus gelada<\/em>): Acoustic Structure and Behavioural Context.\u201d <em>Folia Primatologica<\/em> 55 (3\u20134): 109\u2013132.<\/p>\n<p class=\"import-Normal\">Bell, Sarah A. 2017. \u201cGaldikas, Birute.\u201d In <em>The International Encyclopedia of Primatology, Volume A\u2013G<\/em>, edited by Agust\u00edn Fuentes, 445\u2013446. Malden, MA: John Wiley &amp; Sons.<\/p>\n<p class=\"import-Normal\">Boinski, S. 1992. \u201cOlfactory Communication among Costa Rican Squirrel Monkeys: A Field Study.\u201d <em>Folia Primatologica<\/em> 59 (3): 127\u2013136.<\/p>\n<p class=\"import-Normal\">Cheney, D. L., and R. M. Seyfarth. 1987. \u201cThe Influence of Intergroup Competition on the Survival and Reproduction of Female Vervet Monkeys.\u201d <em>Behavioral Ecology and Sociobiology<\/em> 21 (6): 375\u2013386.<\/p>\n<p class=\"import-Normal\">de Oliveira Terceiro, Francisco Edvaldo, and Judith M. Burkart. 2019. \u201cCooperative Breeding.\u201d In <em>Encyclopedia of Animal Cognition and Behavior<\/em>, edited by Jennifer Vonk and Todd Shackelford, 1\u20136. Edinburg, Scotland: Springer Cham.<\/p>\n<p class=\"import-Normal\">Digby, Leslie J., Stephen F. Ferrari, and Wendy Saltzman. 2011. \u201cCallitrichines: The Role of Competition in Cooperatively Breeding Species.\u201d In <em>Primates in Perspective<\/em>, edited by Christina J. Campbell, August\u00cdn Fuentes, Katherine C. MacKinnon, Simon K. Bearder, and Rebecca M. Stumpf, 91\u201310. 2nd edition. New York: Oxford University Press.<\/p>\n<p class=\"import-Normal\">Fischer, Julia, Kurt Hammerschmidt, Dorothy L. Cheney, and Robert M. Seyfarth. 2008. \u201cAcoustic Features of Female Chacma Baboon Barks.\u201d <em>Ethology<\/em> 107 (1): 33\u201354.<\/p>\n<p class=\"import-Normal\">Jolly, Alison. 1966. <em>Lemur Behavior: A Madagascar Field Study<\/em>. Chicago: University of Chicago Press.<\/p>\n<p class=\"import-Normal\">Krief, Sabrina, Claude Marcel Hladik, and Claudie Haxaire. 2005. \u201cEthnomedicinal and Bioactive Properties of Plants Ingested by Wild Chimpanzees in Uganda.\u201d <em>Journal of Ethnopharmacology<\/em> 110 (1\u20133): 1\u201315.<\/p>\n<p class=\"import-Normal\">Maekawa, Mkio, Annette Lanjouw, Eug\u00e8ne Rutagarama, and Doublas Sharp. 2013. \u201cMountain Gorilla Tourism Generating Wealth and Peace in Post-Conflict Rwanda.\u201d <em>Natural Resources Forum<\/em> 37 (2): 127\u2013137.<\/p>\n<p class=\"import-Normal\">Matsuzawa, Tetsuro. 2015. \u201cSweet-Potato Washing Revisited: 50th Anniversary of the <em>Primates<\/em> Article.\u201d <em>Primates<\/em> 56: 285\u2013287.<\/p>\n<p class=\"import-Normal\">Matsuzawa, Tetsuro. 2018. \u201cHot-Spring Bathing of Wild Monkeys in Shiga-Heights: Origin and Propagation of a Cultural Behavior.\u201d <em>Primates<\/em> 59: 209\u2013213.<\/p>\n<p class=\"import-Normal\">McGrew, W. C. 1998. \u201cCulture in Nonhuman Primates?\u201d <em>Annual Review of Anthropology<\/em> 27: 301\u2013328.<\/p>\n<p class=\"import-Normal\">Mertl-Millhollen, Anne S. 1988. \u201cOlfactory Demarcation of Territorial but Not Home Range Boundaries by <em>Lemur catta<\/em>.\u201d <em>Folia Primatologica<\/em> 50 (3\u20134): 175\u2013187.<\/p>\n<p class=\"import-Normal\">Pinacho-Guendulain, B., and G. Ramos-Fern\u00e1ndez. 2017. \u201cInfluence of Fruit Availability on the Fission-Fusion Dynamics of Spider Monkeys (<em>Ateles geoffroyi<\/em>).\u201d <em>International Journal of Primatology<\/em> 38: 466\u2013484.<\/p>\n<p class=\"import-Normal\">Poirotte, Cl\u00e9mence, Fran\u00e7ois Massol, Ana\u00efs Herbert, Eric Willaume, Pacelle M. Bomo, Peter M. Kappeler, and Marie J. E. Charpentier. 2017. \u201cMandrills Use Olfaction to Socially Avoid Parasitized Conspicifics.\u201d <em>Science Advances<\/em> 3 (4): e160172.<\/p>\n<p class=\"import-Normal\">Rodrigues, Michelle. 2019. \u201cIt\u2019s Time to Stop Lionizing Dian Fossey as a Conservation Hero.\u201d <em>Lady Science<\/em> website, September 20. Accessed December 14, 2022. <a class=\"rId15\" href=\"https:\/\/www.ladyscience.com\/ideas\/time-to-stop-lionizing-dian-fossey-conservation\">https:\/\/www.ladyscience.com\/ideas\/time-to-stop-lionizing-dian-fossey-conservation<\/a>.<\/p>\n<p class=\"import-Normal\">Samuni, Liran, Anna Preis, Tobias Deschner, Catherine Crockford, and Roman M. Wittig. 2018. \u201cReward of Labor Coordination and Hunting Success in Wild Chimpanzees.\u201d <em>Communications Biology<\/em> 1: 138.<\/p>\n<p class=\"import-Normal\">Santana, Sharlene E., Jessica Lynch Alfaro, and Michael E. Alfaro. 2012. \u201cAdaptive Evolution of Facial Colour Patterns in Neotropical Primates.\u201d <em>Proceedings of the Royal Society B: Biological Sciences<\/em> 279 (1736): 2204\u20132211.<\/p>\n<p class=\"import-Normal\">Sanz, Crickette M., David Strait, Crepin Eyana Ayina, Jean Marie Massamba, Thierry Fabrice Ebombi, Severin Ndassoba Kialiema, Delon Ngoteni, et al. 2022. \u201cInterspecific Interactions Between Sympatric Apes.\u201d i<em>Science<\/em> 25 (10): 105059.<\/p>\n<p class=\"import-Normal\">Sch\u00f6n Ybarra, M. A. 1986. \u201cLoud Calls of Adult Male Red Howling Monkeys (<em>Alouatta seniculus<\/em>).\u201d <em>Folia Primatologica<\/em> 47 (4): 204\u2013216.<\/p>\n<p class=\"import-Normal\">Setchell, Joanna M., Tessa Smith, E. Jean Wickings, and Leslie A. Knapp. 2008. \u201cSocial Correlates of Testosterone and Ornamentation in Male Mandrills.\u201d <em>Hormones and Behavior<\/em> 54 (3): 365\u2013372.<\/p>\n<p class=\"import-Normal\">Setchell, Joanna M., E. Jean Wickings, and Leslie A. Knapp. 2006. \u201cSignal Content of Red Facial Coloration in Female Mandrills (<em>Mandrillus sphinx<\/em>).\u201d <em>Proceedings of the Royal Society B: Biological Sciences<\/em><a class=\"rId16\" href=\"https:\/\/paperpile.com\/b\/Gb7Zko\/Lxpl\"> 273 (1599): 2395\u20132400.<\/a><\/p>\n<p class=\"import-Normal\">Seyfarth, R. M., D. L. Cheney, and P. Marler. 1980a. \u201cMonkey Responses to Three Different Alarm Calls: Evidence of Predator Classification and Semantic Communication.\u201d <em>Science<\/em> 210 (4471): 801\u2013803.<\/p>\n<p class=\"import-Normal\">Seyfarth, Robert M., Dorothy L. Cheney, and Peter Marler. 1980b. \u201cVervet Monkey Alarm Calls: Semantic Communication in a Free-Ranging Primate.\u201d <em>Animal Behaviour<\/em> 28 (4): 1070\u20131094.<\/p>\n<p class=\"import-Normal\">Sharma, Goutam, Chan Ram, and Lal Singh Rajpurohit. 2010. \u201cA Case Study of Infantcide After Resident Male Replacement in <em>Semnopithecus entellus<\/em> around Jodhpur (India).\u201d <em>Proceeding of the Zoological Society<\/em> 63 (2): 93\u201398.<\/p>\n<p class=\"import-Normal\">Shirasu, Mika, Satomi Ito, Akihiro Itoigawa, Takashi Hayakawa, Kodzue Kinoshita, Isao Munechika, Hiroo Imai, and Kazushige Touhara. 2020. \u201cKey Male Glandular Odorants Attracting Female Ring-Tailed Lemurs.\u201d <em>Current Biology<\/em> 30 (11): 2131\u20132138.<\/p>\n<p class=\"import-Normal\">Stanford, Craig B. 2017. \u201cGoodall, Jane.\u201d In <em>The International Encyclopedia of Primatology, Volume A\u2013G<\/em>, edited by Agust\u00edn Fuentes, 471\u2013472. Malden, MA: John Wiley &amp; Sons.<\/p>\n<p class=\"import-Normal\">Stewart, Kelly. 2017. \u201cFossey, Dian.\u201d In <em>The International Encyclopedia of Primatology, Volume A\u2013G<\/em>, edited by Agust\u00edn Fuentes, 432\u2013433. Malden, MA: John Wiley &amp; Sons.<\/p>\n<p class=\"import-Normal\">Takeshita, Rafaela S.C., Fred B. Bercovitch, Kodzue Kinoshita, and Michael A. Huffman. 2018. \u201cBeneficial Effect of Hot Spring Bathing on Stress Levels in Japanese Macaques.\u201d <em>Primates<\/em> 59 (3): 215\u2013225.<\/p>\n<p class=\"import-Normal\">Trivers, Robert L. 1972. \u201cParental Investment and Sexual Selection.\u201d In <em>Sexual Selection and the Descent of Man, 1871\u20131971<\/em>, edited by Bernard Campbell, 136\u2013179. Chicago: Aldine.<\/p>\n<p class=\"import-Normal\">Whiten, Andrew. 2011. \u201cThe Scope of Culture in Chimpanzees, Humans and Ancestral Apes.\u201d <em>Philosophical Transactions of the Royal Society of London B: Biological Sciences<\/em> 366 (1567): 997\u20131007.<\/p>\n<p class=\"import-Normal\">Wiens, Frank, and Annette Zitzmann. 2003. \u201cSocial Structure of the Solitary Slow Loris <em>Nycticebus coucang<\/em> (Lorisidae).\u201d <em>Journal of Zoology<\/em> 261 (1): 35\u201346.<\/p>\n<p class=\"import-Normal\">Zuberb\u00fchler, Klaus, David Jenny, and Redouan Bshary. 1999. \u201cThe Predator Deterrence Function of Primate Alarm Calls.\u201d <em>Ethology<\/em> 105 (6): 477\u2013490.<\/p>\n<p class=\"import-Normal\">Zuberb\u00fchler, Klaus, Ronald No\u00eb, and Robert M. Seyfarth. 1997. \u201cDiana Monkey Long-Distance Calls: Messages for Conspecifics and Predators.\u201d <em>Animal Behaviour<\/em> 53 (3): 589\u2013604.<\/p>\n<h2 class=\"import-Normal\">Acknowledgements<\/h2>\n<p class=\"import-Normal\">The author is grateful to the editors for the opportunity to contribute to this open-source textbook. She thanks Dr. Stephanie Etting for her encouragement and support during the revision of this chapter. Her suggestions, along with comments made by two anonymous reviewers on an earlier draft of this chapter, improved the final version considerably. Finally, she thanks all the primatologists who came before her, especially her advisor, Lynne A. Isbell, for their tireless efforts to understand the behavior and ecology of the living primates. Without their work, this chapter would not have been possible.<\/p>\n<\/div>\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_1683_1713\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1713\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<div class=\"textbox\">\n<p>Learn more about <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/about-the-authors\/\">authors and editors<\/a>,<a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/student-contributors\/\"> Hess' student contributions<\/a>, and <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/versioning-history\/\">versioning history<\/a>.<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\">Learning Objectives<\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li class=\"import-Normal\">Describe the variables that affect primate diets.<\/li>\n<li class=\"import-Normal\">Explain how primates interact with other organisms in their environment.<\/li>\n<li class=\"import-Normal\">Discuss why primates live in groups, types of primate groups, and components of their social systems.<\/li>\n<li class=\"import-Normal\">Describe the reproductive strategies of males and females.<\/li>\n<li class=\"import-Normal\">Explain the ways in which primates communicate.<\/li>\n<li class=\"import-Normal\">Discuss the evidence for primate cultural traditions.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p class=\"import-Normal\">Nonhuman primates (hereafter, 'primates') are a fascinating group of animals, whose similarity to humans can be striking. Because of this similarity, studying primates helps anthropologists to gain insight into how our human ancestors may have behaved. It also allows us to better understand our own behaviour through <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_928\">comparison<\/a><\/strong> (examining similarities and differences) with other primates as well as by comparing different species of primates to one another. In this way, studying primates helps anthropologists comprehend humanity from a biological perspective, which contributes to anthropology\u2019s commitment to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_930\">holism,<\/a><\/strong> the idea that the parts of a system interconnect and interact to make up the whole.<\/p>\n<figure style=\"width: 242px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2023\/05\/image3.png\" alt=\"A person using binoculars to look at monkeys.\" width=\"242\" height=\"354\" \/><figcaption class=\"wp-caption-text\">Figure 7.1: The author observing patas monkeys from a distance in Laikipia, Kenya. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-6\/\">Karin Enstam Jaffe observing patas monkeys in Laikipia, Kenya (Figure 6.5)<\/a> by Rebecca Chancellor is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_932\">Ethology<\/a> <\/strong>is the study of animal behaviour, while <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_934\">primatology <\/a><\/strong>is the study of primate behaviour. People who study primates are called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_936\">primatologists<\/a><\/strong>. Research on primates can be conducted in the field (i.e., on wild primates) or in captivity (i.e., zoos) and may or may not involve experiments, such as playing recorded alarm calls to see how individuals react. Unlike some other Science, Technology, Engineering, and Math (STEM) fields, primatology has a long history of research conducted by women (see \u201cSpecial Topic: Women in Primatology\u201d). Primatologists come from many different disciplines, have diverse backgrounds, and study primates for different reasons. Biologists study primates as examples of evolutionary theories like natural selection, and to understand behaviours as <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_938\">adaptations<\/a><\/strong>, or traits with a function that increases <strong>fitness<\/strong>, i.e. an individual\u2019s survival and\/or reproduction. Primate intelligence is of interest to psychologists who want to learn more about deception or cooperation and to linguists interested in the principles of communication and language. Ecologists consider how primates interact with the habitats they occupy, and conservationists examine how primates are affected by deforestation, poaching, or illegal animal trade (see Appendix B: Primate Conservation for more information on these topics). Biological anthropologists, like myself (Figure 7.1), who study primates are interested in learning about their social complexity, and ecological and behavioural variation, to better understand the biological basis of human behaviour. And, similar to biologists, we also explore how primate behaviour is adaptive and contributes to individual fitness. Like other sciences, primatology is only as strong as its researchers, methods, and theories, and the field has benefitted recently from efforts to increase diversity and reckon with its colonialist past, as discussed below in \u201cSpecial Topic: Women in Primatology.\u201d<\/p>\n<p class=\"import-Normal\">Humans share many traits in common with primates. As you learned in Chapter 5, some of these traits are similar due to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_944\">homology<\/a><\/strong>, traits both species inherited from a common primate ancestor. For example, like most other primates, humans are social animals who live in groups. Group living did not evolve independently in humans and other primates. Rather, group living is a trait that evolved in a primate ancestor, and because it benefited survival, it was retained in the species\u2019 <strong>descendants<\/strong> (or the species that come after the ancestor species). In contrast, humans and other primates can have similar traits that evolved independently, which is called <strong>analogy<\/strong>. For example, both humans and Japanese macaques (<em>Macaca fuscata<\/em>) use natural hot springs (Figures 7.2a &amp; b). Research on these monkeys indicates that sitting in hot springs reduces stress and helps keep them warm, much as it does for humans (Takeshita Et al. 2018). But this behaviour is not the result of humans and Japanese macaques having a shared ancestor who used hot springs. Rather, the behaviour arose independently in two species that both occupy northerly environments and adapted to cold climates using a similar behaviour. Studying the homologous traits we share with other primates, like living in groups, helps us develop hypotheses about human behaviours as adaptations, which in turn helps us develop models for the behaviour of our human ancestors. Studying analogous traits, like hot springs use, allows us to better understand the effects of ecological variables on morphology and behaviour of both primates and humans, living and extinct.<\/p>\n<figure id=\"attachment_191-3\" aria-describedby=\"caption-attachment-191-3\" style=\"width: 2161px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-189 size-full\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.2.jpg\" alt=\"Left, a man in a hot spring. Right, monkeys in a hot spring.\" width=\"2161\" height=\"803\" \/><figcaption id=\"caption-attachment-191-3\" class=\"wp-caption-text\">Figure 7.2a\/b: Both humans (left) and Japanese macaques (right) use natural hot springs to reduce stress and relax. This similar trait arose independently in the two species, making it a good example of analogy. Credit: 7.2a. <a href=\"https:\/\/pixabay.com\/photos\/hot-spring-landscape-man-mountain-1846721\/\">Hot Spring Landscape<\/a> by <a href=\"https:\/\/pixabay.com\/users\/pexels-2286921\/?utm_source=link-attribution&amp;utm_medium=referral&amp;utm_campaign=image&amp;utm_content=1846721\">Pexels<\/a> has been modified (cropped) and 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>. 7.2b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Jigokudani_hotspring_in_Nagano_Japan_001.jpg\">Jigokudani hotspring in Nagano Japan 001<\/a> by Yosemite 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<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Women in Primatology<\/h2>\n<p class=\"import-Normal\">While many STEM fields have traditionally been, and continue to be, dominated by men, primatology has a long history of significant research conducted by women. This is due, in part, to the fact that three of the most well-known primatologists are women. In the early 1960s, British paleoanthropologist Louis Leakey (discussed in Chapters 9 and 10) was looking for students to study the great apes in hopes of shedding light on the behaviours of our early ancestors. He chose Jane Goodall (Figure 7.3a) to study chimpanzees (<em>Pan troglodytes<\/em>), Birute Galdikas (Figure 7.3b) to study Bornean orangutans (<em>Pongo pygmaeus<\/em>), and Dian Fossey (Figure 7.3c) to study mountain gorillas (<em>Gorilla<\/em><em> beringei beringei<\/em>). The work of these three women, sometimes referred to as Leakey\u2019s \u201cTrimates,\u201d has transformed our understanding of ape (and primate) behaviour.<\/p>\n<figure id=\"attachment_190-2\" aria-describedby=\"caption-attachment-190-2\" style=\"width: 626px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-190\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.3-1.jpg\" alt=\"Jane Goodall, Birute Galdikas, and Dian Fossey.\" width=\"626\" height=\"209\" \/><figcaption id=\"caption-attachment-190-2\" class=\"wp-caption-text\">Figure 7.3a-c: Louis Leakey\u2019s \u201cTrimates\u201d (left to right): a. Jane Goodall\u2019s research on the Gombe chimpanzees spans over half a century; b. Birute Galdikas\u2019s research and rescue work on behalf of orangutans spans 40 years; c. Dian Fossey studied mountain gorillas in Rwanda for almost 20 years, until her murder in 1985. Credit: a. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Jane_Goodall_HK.jpg\">Jane Goodall HK<\/a> by Jeekc has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\"> CC BY-SA 3.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Dr_Birute_Galdikas.jpg\">Dr Birute Galdikas<\/a> by<a href=\"https:\/\/www.flickr.com\/photos\/sfupamr\/\"> Simon Fraser University - University Communications<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\"> CC BY 2.0 License<\/a>. c. <a href=\"https:\/\/www.flickr.com\/photos\/mary-lynn\/2925879356\">US-223658 Dian Fossey<\/a> by<a href=\"https:\/\/www.flickr.com\/photos\/mary-lynn\/2925879356\/\"> Mary-Lynn<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\"> CC BY 2.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Arriving at the Gombe Stream Reserve in Tanzania in 1960, Jane Goodall was one of the first scientists to conduct a long-term study of wild nonhuman primates. Before then, most studies lasted less than a year and were often zoo-based. By 1961, she had made two astounding observations that forced us to reconsider what differentiates humans from the rest of the primate order. She observed chimpanzees eating a colobus monkey, the first reported evidence of meat eating in our closest relatives; later observing them hunting and sharing meat. And she discovered that chimpanzees make and use tools by stripping leaves off twigs to \u201cfish\u201d for termites. Her work, spanning several decades, has produced long-term data on chimpanzee mating strategies, mother-infant bonds, and aggression. In the mid-1980s, Goodall transitioned from field researcher to conservationist and activist, advocating for the humane use of nonhuman animals (Stanford 2017).<\/p>\n<p class=\"import-Normal\">Birute Galdikas began her study of orangutans in Kalimantan, Borneo, in 1971. Hers was the first long-term study conducted on the Bornean orangutan. Galdikas and her colleagues have collected over 150,000 hours of observational data, focusing on the life histories of individual orangutans. While conducting behavioural research, Galdikas discovered that the pet trade and habitat loss were adversely affecting the orangutan population. Eventually, Galdikas\u2019s conservation efforts began to extend beyond advocacy and into rehabilitation and forest preservation (Bell 2017). If you would like to learn more about primate conservation efforts, please see Appendix B: Primate Conservation.<\/p>\n<p class=\"import-Normal\">In 1967, Dian Fossey began her long-term study of mountain gorillas and founded the Karisoke Research Centre in Rwanda. Her and her colleagues\u2019 research, over several decades, revealed much about gorilla social behaviour, ecology, and life history. Her efforts also led to the development of mountain gorilla conservation programs. However, she was a controversial figure, as discussed below. Fossey was murdered in December 1985; the case remains unsolved (Stewart 2017).<\/p>\n<h3 class=\"import-Normal\"><strong>Decolonizing Primatology<\/strong><\/h3>\n<p class=\"import-Normal\">Recently, the movement to <strong>decolonize<\/strong> primatology, by understanding and highlighting the theories and research of non-Western individuals and perspectives, has gathered steam. This movement draws attention to the maltreatment of local people by Western primatologists. For example, Michelle Rodrigues (2019) argues that it's time we stop focusing on the scientific and conservation contributions of Dian Fossey and acknowledge that her \"active conservation\" techniques included kidnapping and torturing local Rwandans who were known as, or suspected to be, gorilla poachers. Rodrigues (2019) argues:<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 36pt;text-indent: 0pt\">The image of Fossey, a white American woman, whipping and torturing black African poachers is evocative of the behaviour of white slaveholders in the American South. It is appalling enough to think of that behaviour occurring in the 1850s; there is no way we can explain Fossey\u2019s behaviour in the 1970s as the product of \u201ca different time.\u201d Yet, almost three decades later, the romantic notion of a noble martyr who died for her devotion to gorillas prevails, and these terrifying actions are often described as simply unorthodox methods. Perhaps these truths are softened due to fears that the reality of this legacy would harm gorilla conservation efforts. But memorializing her as a martyr and patron saint of gorilla conservation demands that we forget the cruel acts she advocated for and performed.<\/p>\n<p class=\"import-Normal\">Further, Louis Leakey\u2019s installment of Goodall, Galdikas, and Fossey to study chimpanzees, orangutans, and mountain gorillas, respectively, is itself viewed as recapitulating the colonial legacy in Africa and Asia. Given that Leakey was the offspring of British missionaries, Rodrigues (2019) argues, it is no accident that he was willing to mentor British and American women, while overlooking women from Africa and Asia as potential researchers. This leads us to another level of the decolonizing movement, which aims to highlight the research of non-Western primatologists, particularly those living in what primatologists refer to as \u201chabitat countries\u201d that are home to living primates. As you will see in this chapter, scientists from diverse backgrounds are active contributors to exciting research on primates around the world.<\/p>\n<\/div>\n<h2 class=\"import-Normal\">Ecology<\/h2>\n<p class=\"import-Normal\">The more than 600 species and subspecies of living primates are highly diverse in their dietary preferences and the habitats they occupy. In this section we\u2019ll briefly discuss aspects of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_950\">ecology<\/a><\/strong>, or the relationship between organisms and their physical surroundings, that impact a primate\u2019s life, the foods they eat, and the other species with whom they interact.<\/p>\n<h3 class=\"import-Normal\"><strong>Primate Diets<\/strong><\/h3>\n<p class=\"import-Normal\">Diet may be the most important variable influencing variation in primate morphology, behaviour, and ecology. Most primates are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_952\">omnivores<\/a><\/strong> who ingest a variety of foods in order to obtain appropriate levels of protein, carbohydrates, fats, and fluids, but one type of food often makes up the majority of each species\u2019 diet. You learned about the dental and digestive adaptations of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_954\">frugivores<\/a><\/strong> (who feed primarily on fruit), <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_956\">folivores<\/a><\/strong> (whose diet consists mostly of leaves), and <strong>insectivores <\/strong>(who eat mainly insects) in Chapter 6, so we will not discuss them again here.<\/p>\n<h4 class=\"import-Normal\"><em>Body Size and Diet<\/em><\/h4>\n<figure id=\"attachment_191-4\" aria-describedby=\"caption-attachment-191-4\" style=\"width: 559px\" class=\"wp-caption alignright\"><img class=\"wp-image-191\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.4-1.jpg\" alt=\"A tarsier eats a grasshopper. A gorilla eats leaves.\" width=\"559\" height=\"237\" \/><figcaption id=\"caption-attachment-191-4\" class=\"wp-caption-text\">Figure 7.4a-b: Primates eat different types of food. Small primates, like the spectral tarsier (left), eat mostly insects while large primates, like the mountain gorilla (right), eat mostly leaves. Credit: a. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Spectral_Tarsier_Tarsius_tarsier_(7911549768).jpg\">Spectral Tarsier Tarsius tarsier (7911549768)<\/a> by<a href=\"https:\/\/www.flickr.com\/people\/65695019@N07\"> Bernard DUPONT<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/legalcode\"> CC BY-SA 2.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Mountain_gorilla_(Gorilla_beringei_beringei)_eating.jpg\">Mountain gorilla (Gorilla beringei beringei) eating<\/a> by<a href=\"https:\/\/www.sharpphotography.co.uk\/\"> Charles J Sharp<\/a> (creator QS:P170,Q54800218) has been modified (cropped) 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>Insects are a high-quality food, full of easily digestible protein and high in calories that meet most of a primate\u2019s dietary needs. Although all primates will eat insects if they come upon them, those species that rely most heavily on insects tend to be the smallest. Why? Because larger primates simply cannot capture and consume enough insects every day to survive. Because of their small size (less than 150 g), spectral tarsiers (<em>Tarsius spectrum<\/em>) have a fast <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_960\">metabolism<\/a><\/strong>, which means they turn food to energy quickly, but they do not need to consume large amounts of food each day. It does not matter to a spectral tarsier that a grasshopper only weighs 300 mg, because the tarsier (<em>Tarsius<\/em>) itself is so small that one grasshopper is a good-size meal (Figure 7.4a). That same grasshopper is not even a snack for an adult male mountain gorilla (<em>Gorilla beringei beringei<\/em>), who may weigh up to 200 kg. Fortunately for gorillas (<em>Gorilla)<\/em>, their large body size means they have a slow metabolism, converting food into energy much more slowly, so they can eat lower quality food that takes longer to digest, provided there is a lot of it. For gorillas, leaves, which are hard to digest but plentiful, fit the bill (Figure 7.4b). Most medium-sized primates are highly frugivorous, and supplement their fruit based diet in ways that correspond with their size: Smaller frugivores tend to supplement with insects, while larger frugivores tend to supplement with leaves.<\/p>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<h4 class=\"import-Normal\"><em>Food Abundance and Distribution<\/em><\/h4>\n<p class=\"import-Normal\">Nutrients are not the only dietary considerations primates must make. They must also ensure that they consume more calories than they use. The abundance and distribution of food affect energy expenditure and calorie intake because they determine how far animals must travel in search of food and how much they must compete to obtain it. <strong>Abundance <\/strong>refers to how much food is available in a given area while <strong>distribution<\/strong> refers to how food is spread out. In terms of abundance, food is either plentiful or scarce (Figure 7.5a\u2013b). Food is distributed in one of three ways: uniformly (Figure 7.6a), in clumps (Figure 7.6b), or randomly (Figure 7.6c). In general, higher-quality foods, like fruit and insects, are less abundant and have patchier distributions than lower-quality foods, like leaves. Primates who eat fruit or insects usually have to travel farther to find food and burn more calories in the process. Abundance and distribution of food is another reason why larger primates tend to rely more heavily on leaves than either fruit or insects.<\/p>\n<figure style=\"width: 619px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image2-2.png\" alt=\"Two squares with different amounts of dots. \" width=\"619\" height=\"286\" \/><figcaption class=\"wp-caption-text\">Figure 7.5a-b: Two types of food abundance. Food is plentiful when there is a lot of it in a given area (left). Food is scarce when there is not very much of it in a given area (right). Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-6\/\">Food abundance and food scarcity (Figure 6.7)<\/a> by Karin Enstam Jaffe 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<figure style=\"width: 690px\" class=\"wp-caption aligncenter\"><img style=\"font-size: 1em\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-2.png\" alt=\"Three squares with dots in different formations. \" width=\"690\" height=\"214\" \/><figcaption class=\"wp-caption-text\">Figure 7.6a-c: Three types of food distribution. a. Food has a uniform distribution when it is spread out evenly in the environment. b. Food has a clumped distribution when it is found in patches. c. Food is randomly distributed when it has neither uniform nor clumped distribution. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-6\/\">Food distribution patterns (Figure 6.8)<\/a> by Karin Enstam Jaffe 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<h3 class=\"import-Normal\"><strong>Community Ecology<\/strong><\/h3>\n<p class=\"import-Normal\">Primates are members of broader ecological communities composed of other species, including other primates, predators, parasites, and even humans. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_966\">Community ecology<\/a><\/strong> deals with the relationships and interactions between different organisms that occupy the same habitat. Interactions with <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_968\">conspecifics<\/a> <\/strong>(members of the same species) and <strong>heterospecifics<\/strong> (members of different species) are critical aspects of ecological communities. Some habitats support highly diverse <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_972\">primate communities<\/a><\/strong> consisting of 10 or more primate species. How can so many primate species occupy the same area and avoid competition? In most cases, the primate species that live together occupy different <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_974\">niches<\/a><\/strong>, which means they do not meet their needs for food and shelter in the exact same way. Two species can avoid competition by eating different kinds of food, living at different levels of a forest, or even searching for food at different times of day. Because tropical rainforests, like Manu National Park in Peru, are highly variable, with many habitats and many sources of food and shelter, there are many different niches for multiple species to exploit, and large primate communities can result (Figure 7.7).<\/p>\n<figure style=\"width: 710px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5-1-1.png\" alt=\"Eight primate species.\" width=\"710\" height=\"550\" \/><figcaption class=\"wp-caption-text\">Figure 7.7: Eight of the 14 primate species in Manu National Park, Peru. Top row, left to right: Goeldi\u2019s marmoset (Callimico goeldi), Rio Tapaj\u00f3s saki (Pithecia irrorata), tufted capuchin (Sapajus apella); middle row, left to right: emperor tamarin (Saguinus imperator), black-headed night monkey (Aotus nigriceps), Bolivian red howler (Alouatta sara); bottom row, left to right: black-capped squirrel monkey (Saimiri boliviensis), Peruvian spider monkey (Ateles chamek). Credit: Primate species in Manu National Park original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Jaffe 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:\/\/www.flickr.com\/photos\/31223088@N08\/5582747190\/\">Tamarin Baby\/Goeldi\u2019s Monkey<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/31223088@N08\/\">stefan_fotos<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\">CC BY 2.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Pithecia_irrorata_-Brazil-8b.jpg\">Pithecia irrorata -Brazil-8b<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/9092428@N04\">Ana_Cotta<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/deed.en\">CC BY 2.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Tufted_capuchin_on_a_branch_in_Singapore.jpg\">Tufted Capuchin on a Branch in Singapor<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Basile_Morin\">Basile_Morin<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Tamarin_portrait.JPG\">Tamarin Portrait<\/a> by <a href=\"https:\/\/sites.google.com\/site\/thebrockeninglory\/?pli=1\">Brocken Inaglory<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/deed.en\">CC BY-SA 3.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Aotus_nigriceps_1.jpg\">Aotus nigriceps 1<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/dusantos_bh\/\">DuSantos<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/deed.en\">CC BY 2.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Alouatta_sara_%28Bolivian_red_howler%29.jpg\">Aloutta sara (Bolivian Red Howler)<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/nacho_dayz\/\">Raul Ignacio<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\">CC BY-SA 2.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Black-capped_squirrel_monkey_%28Chalalan%29.jpg\">Black-Capped Squirrel Monkey (Chalalan)<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Rodrigo_Mariaca\">Rodrigo Mariaca<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0 License<\/a>; <a href=\"https:\/\/www.flickr.com\/photos\/eye1\/3185562151\/\">Maquisapa (Spider Monkey)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/eye1\/\">Ivan Mlinaric<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\">CC BY 2.0 License<\/a>].<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Competitive Interactions<\/em><\/h4>\n<p class=\"import-Normal\">Although species living in the same location often occupy different niches to avoid competition, when a resource that is important for survival or reproduction is scarce, individuals will compete to obtain that resource. This is a central tenet of Charles Darwin\u2019s theory of evolution by natural selection (see Chapter 2 &amp; 3). Competition between primates takes two forms: Individuals engage in <strong>direct competition<\/strong>, which involves physical interaction between individuals (such as fighting), over resources that are large and worth defending (fruit is a good example of a food resource over which primates will fight). Individuals engage in <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_978\">indirect competition<\/a><\/strong>, in which there is no physical interaction between individuals, when a resource is small. Primates often engage in indirect competition for insects, like grasshoppers, that are eaten quickly, often before another individual arrives on the scene. Primates may engage in direct and\/or indirect competition with members of their own group, with members of other groups of conspecifics, or with heterospecifics.<\/p>\n<h4 class=\"import-Normal\"><em>Predator-Prey Interactions<\/em><\/h4>\n<p class=\"import-Normal\">The plants and animals that primates eat are an important part of their ecological community. In addition to insects, many primates incorporate some <strong>vertebrate<\/strong> (animals with an internal spinal column or backbone) prey into their diet. Often, predation by primates is opportunistic, occurring because the prey happens to be in the right place at the right time. I\u2019ve observed vervets (<em>Chlorocebus pygerythrus<\/em>) opportunistically killing lizards by smashing them against a rock or tree trunk and eating them. More rarely, hunting is deliberate and cooperative. In some chimpanzee (<em>Pan troglodytes<\/em>) populations, hunts involve multiple individuals, each of whom plays a specific role and is rewarded afterward with a share of the prey that has been captured (Samuni Et al. 2018).<\/p>\n<p class=\"import-Normal\">All primates are susceptible to predation by mammalian <strong>carnivores <\/strong>(animals whose diet consists primarily of animal tissue (e.g., Figure 7.8a), reptiles (e.g., Figure 7.8b), or birds of prey (e.g., Figure 7.8c). Although the specific predators found in an ecological community differ based on geography, smaller primates always fall prey to a wider range of predators. Because predators are diverse in their hunting tactics, primates have evolved a wide range of tactics to avoid or escape them. We will discuss some of these behavioural adaptations later in this chapter in the section titled \u201cWhy Do Primates Live in Groups?.\u201d<\/p>\n<p>&nbsp;<\/p>\n<\/div>\n<figure id=\"attachment_210-8\" aria-describedby=\"caption-attachment-210-8\" style=\"width: 765px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-195\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.8-scaled-1.jpg\" alt=\"A leopard, python, and harpy eagle.\" width=\"765\" height=\"229\" \/><figcaption id=\"caption-attachment-210-8\" class=\"wp-caption-text\">Figure 7.8a-c: Examples of primate predators: the Indian leopard (Panthera fusca) is an example of a mammalian carnivore (top left), the South African python (Python natalensis) is an example of a reptilian predator (bottom left), and the harpy eagle (Harpia harpyja) of Central and South America is an example of a bird of prey (right). Credit: a. <a href=\"https:\/\/www.flickr.com\/photos\/srikaanth-sekar\/9814267145\/\">Leopard<\/a> by<a href=\"https:\/\/www.flickr.com\/people\/srikaanth-sekar\/\"> Srikaanth Sekar<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/\"> CC BY-SA 2.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Python_natalensis_G._J._Alexander.JPG\">Python natalensis G. J. Alexander<\/a> by Graham J. Alexander, University of the Witwatersrand, USGS, is in the<a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\"> public domain<\/a>. c. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Harpy_Eagle_clutching_captured_bird_-_Itirapina_Reserve.jpg\">Harpy Eagle clutching captured bird - Itirapina Reserve<\/a> by Jonathan Wilkins has been modified (cropped) and 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<div class=\"__UNKNOWN__\">\n<h4 class=\"import-Normal\"><em>Mutualistic Interactions<\/em><\/h4>\n<p class=\"import-Normal\">So far, we've discussed competitive and predator-prey interactions in primate communities. But there are some interactions (between different primate species and between primates and other species) that are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_984\">mutualistic<\/a><\/strong>, which is when organisms of different species work together, each benefiting from the interaction or relationship. One example is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_986\">seed dispersal<\/a><\/strong>, which is the process by which seeds move away from the plant that produced them in preparation for germination and becoming a new plant. When seeds are dispersed by animals, like primates, it is an example of mutualism. The primate eats the fruit of a plant, which provides nutrients for its body, and in the process ingests the plant\u2019s seeds. Later, it deposits the seeds at another location as a pile of fertilizer.<\/p>\n<p class=\"import-Normal\">Another example of mutualism is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_988\">polyspecific associations<\/a><\/strong>, which are associations between two or more different species that are maintained by behavioural changes by at least one of the species. While some associations are short in duration, others are semi-permanent. The mutualistic benefits of polyspecific associations include one species gaining access to food that would otherwise have been inaccessible or being alerted to the presence of predators that they would not have not have known were present otherwise. In some cases, individuals seem to recognize and seek out specific members of another species. Twenty years of observations on chimpanzees and Western lowland gorillas (<em>Gorilla gorilla gorilla<\/em>) in the Republic of Congo has revealed social ties (some might call them friendships) between individual chimpanzees and gorillas that last for years and occur in a variety of social contexts, including play (Sanz Et al. 2022).<\/p>\n<h4 class=\"import-Normal\"><em>Parasite-Host Interactions<\/em><\/h4>\n<p class=\"import-Normal\">Primates are hosts for a variety of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_990\">parasites<\/a><\/strong>, which are organisms that live in or on another organism (the host). Parasites come in many forms and pose varying levels of danger to the host. Blood parasites cause diseases like yellow fever and malaria. Skin parasites include fleas and ticks, which feed on the host\u2019s blood, and botflies, which lay eggs in the host\u2019s flesh. Bot fly larvae feed on the host\u2019s flesh as they develop and eventually (if not removed) break through the skin at maturity. Gut parasites, like tapeworms, get into the intestines and feed off of the food that is being digested by the host. Because most primates live in groups (see the \u201cPrimate Societies\u201d section of this chapter), the tendency for <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_992\">social transmission<\/a><\/strong> of parasites, or the transfer of parasites from one individual to another, is high. Primates have evolved mechanisms to avoid parasite infection, including switching sleeping and feeding sites so as to avoid parasites. Mandrills (<em>Mandrillus sphinx<\/em>) have been shown to avoid grooming infected conspecifics as well as to avoid their feces, which smell different than the feces of individuals who are not infected with parasites (Poirotte Et al. 2017). Other primates, including chimpanzees, appear to self-medicate when infected with parasites by ingesting plants that have antiparasitic properties (Krief Et al. 2005).<\/p>\n<h4 class=\"import-Normal\"><em>Human-Primate Interactions<\/em><\/h4>\n<p class=\"import-Normal\">Humans are part of many primate communities and our relationship with our closest relatives is often complicated. In some areas, humans hunt primates for their meat or as trophies, or so they can sell the infants as pets. As the human population increases in size, our demand for natural resources, like wood to build houses or land on which to grow food, also increases, often at the expense of pristine primate (and other animal) habitat. As their natural habitat shrinks, primates search for food in areas occupied by humans and may be shot as crop-raiding pests. While deforestation, hunting, and the pet trade are examples of ways in which humans negatively affect the lives of primates, some human-primate interactions are beneficial. In some parts of the world primates are central to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_994\">ecotourism<\/a><\/strong>, which focuses on nature-based attractions to educate tourists and uses economically and ecologically sustainable practices. Perhaps one of the greatest success stories of ecotourism involves the mountain gorillas of Rwanda (see Figure 7.4b). After internal conflict plagued Rwanda during the 1990s, the Virunga Mountains area developed gorilla-based tourism to aid in socioeconomic development and to bring stability to the region. This process not only helped to increase mountain gorilla populations but was also able to generate enough income to cover the operation costs of three national parks and provide income and other benefits to people living in the area (Maekawa Et al. 2013). You can learn more about human-primate interactions in Appendix B: Primate Conservation.<\/p>\n<h2 class=\"import-Normal\">Primate Societies<\/h2>\n<p class=\"import-Normal\">Unlike many other animals, primates are highly social and many live in stable groups consisting of adult males and females, even outside the<strong> <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_996\">breeding season<\/a><\/strong>, when females are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_998\">receptive<\/a><\/strong> and available for mating because they are not pregnant or nursing. Indeed, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1000\">sociality<\/a><\/strong>, or the tendency to form social groups, is a key behavioural adaptation of the order primates (see Chapter 6). This has led primatologists to ask two questions: \u201cWhy do primates live in groups?\u201d and \u201cWhat types of groups do primates live in?\u201d<\/p>\n<h3 class=\"import-Normal\"><strong>Why Do Primates Live in Groups?<\/strong><\/h3>\n<p class=\"import-Normal\">Primates live in groups when the benefits of doing so exceed the costs. Although there are many potential benefits to group living, enhanced feeding competition and predator avoidance are important benefits for many group living primates. When primates feed on high-quality, scarce food (like fruit), larger groups are more successful in competition with other groups. For example, in a long-term study of vervets in Kenya\u2019s Amboseli National Park, larger vervet groups had larger and better <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1002\">home ranges<\/a><\/strong>, which is the area in which a group regularly moves around as it performs its daily activities, including searching for food and water. Females in larger groups had higher average infant and female survival rates than the smallest group. Because pregnancy and nursing are energetically expensive for females, female <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1004\">reproductive success<\/a><\/strong>, or genetic contribution to future generations (measured by the number of offspring produced), is limited by access to food. Although living in a group means females compete with members of their own group for food, the benefits of being a member of a larger vervet group outweigh the costs (Cheney &amp; Seyfarth 1987).<\/p>\n<p class=\"import-Normal\">However, because they contain more individuals, larger groups are more likely to attract the attention of predators compared to smaller groups. This is one of the reasons that primates who rely on <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1006\">crypsis<\/a><\/strong>, or the ability to avoid detection by others, including predators, are often <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1008\">solitary<\/a> <\/strong>(the term used to describe individuals who do not live together with other members of their species) and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1010\">nocturnal<\/a><\/strong>, or active at night. If an animal is already hard to see because it is active at night, then moving quietly in small groups is a good strategy to avoid detection by predators. The slow loris (<em>Nycticebus coucang<\/em>) of Southeast Asia is a good example of this strategy (Figure 7.9a). Nocturnal and solitary, the slow loris moves slowly and quietly as its primary strategy to avoid detection (Wiens &amp; Zitzmann 2003). In contrast, primates who live in large groups and are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1012\">diurnal<\/a><\/strong>, or active during the day (like gelada baboons [<em>Theropithecus gelada<\/em>]; Figure 7.9b) cannot avoid detection by predators. Instead, group-living primates rely on behaviours that alert others to the presence of danger and\/or deter predators, including shared <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1014\">vigilance<\/a><\/strong> (watchful behaviour to detect potential danger), <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1016\">mobbing<\/a><\/strong> (the act of cooperatively attacking or harassing a predator), and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1018\">alarm calling<\/a><\/strong> (vocalizations emitted by social animals in response to danger). We will discuss alarm calls in the <em>Communication<\/em> section.<\/p>\n<figure id=\"attachment_196-2\" aria-describedby=\"caption-attachment-196-2\" style=\"width: 761px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-196\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.9.jpg\" alt=\"A slow loris. A group of gelada baboons.\" width=\"761\" height=\"269\" \/><figcaption id=\"caption-attachment-196-2\" class=\"wp-caption-text\">Figure 7.9a\/b: Some primates, like the slow loris (left), are solitary and spend most of their time alone. However, most primates, like the gelada baboon (right), live in groups of varying sizes. Credit: a. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Slow_Loris.jpg\">Slow Loris<\/a> by Jmiksanek is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\"> CC BY-SA 3.0 License<\/a>. b. <a href=\"https:\/\/www.flickr.com\/photos\/39997856@N03\/7588490544\">Field of baboons<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/39997856@N03\/\">mariusz kluzniak<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/2.0\/\">CC BY-NC-ND 2.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>What Types of Groups Do Primates Live In? <\/strong><\/h3>\n<p class=\"import-Normal\">Primates vary with regard to the types of groups in which they live. A <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1020\">social system<\/a> <\/strong>describes a set of social interactions and behaviours that is typical for a species. The components that make up a species\u2019 social system include:<\/p>\n<ul>\n<li class=\"import-Normal\">Group size, which refers to the number of individuals that typically live together. Primate group size can be highly variable, ranging from one or a few individuals, to a few dozen, upward to several hundred individuals.<\/li>\n<li class=\"import-Normal\">Group composition describes group membership in terms of age class (e.g., adult, juvenile, infant) and sex. In some primates, groups consist of a mother and her dependent offspring while in others, one adult male lives long-term with one adult female and their dependent offspring. In other species, one or more adult males live with multiple females and their offspring.<\/li>\n<li class=\"import-Normal\">A species\u2019 <strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1022\">mating system<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"> refers to which male(s) and female(s) mate. The terms that describe a mating system (e.g., <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1024\">polygyny<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">, in which one male mates with multiple females) are sometimes used to describe a primate species\u2019 social system, but a mating system is one component of the species\u2019 social system. For example, two species might both have polygynous mating systems, but in one species, the group is composed of one male and multiple females, while members of the other species live as solitary individuals.<\/span><\/li>\n<li class=\"import-Normal\"><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1026\">Ranging behaviour<\/a> <\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">refers to the way in which animals move about their environment. Most primate species have a home range, where they perform their daily activities. Some primates defend a <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1028\">territory<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"> which is the part of the home range that the group actively guards in an attempt to keep out conspecifics.<\/span><\/li>\n<li class=\"import-Normal\"><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1030\">Dispersal<\/a> <\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">patterns describe which sex moves to a new group to reproduce. In most primate species, males disperse because the benefits of dispersal, including increased access to mates and reduced competition from other males, outweigh the costs of migrating into a new group, which often comes with aggression from current group members. For many female primates, the opposite is true: females usually benefit from remaining <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1032\">philopatric<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">, or in the group of their birth. This allows them to maintain strong alliances with female relatives, which helps them compete successfully against other groups for food. In solitary species, offspring of both sexes leave their mother\u2019s home range and become solitary. If this did not happen, the species would not be solitary. Even though both sexes disperse in solitary species, males usually disperse farther than females.<\/span><\/li>\n<li class=\"import-Normal\">Social interactions describe the ways in which individuals interact with members of their own and other groups of conspecifics. <strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1034\">Affiliative<\/a> <\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">(i.e., friendly or nonaggressive) behaviours include <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1036\">grooming<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"> (picking through the fur of another individual), playing, or <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1038\">coalitions<\/a> <\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">(temporary alliances between individuals). <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1040\">Agonistic<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"> (i.e., aggressive) behaviours include fighting over food or fighting over access to mates. In groups that contain multiple adult individuals of the same sex, it is common to have a <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1042\">dominance hierarchy<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">, or a group of individuals that can be ranked according to their relative amount of power over others in the hierarchy. Initially, dominance hierarchies are established through the outcome of conflicts. Individuals who lose conflicts with others are <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1044\">subordinate<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"> (or low rank) to those who win them. Those who win conflicts are <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_744\">dominant<\/a> <\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">(or high rank). Dominant individuals gain access to resources, like food or mates, before subordinates. Once a hierarchy is established, agonism decreases because everyone \u201cknows their place.\u201d<\/span><\/li>\n<\/ul>\n<p class=\"import-Normal\">The main types of primate social systems are as follows: solitary; single-male, single-female; single-male, multi-female; multi-male, multi-female; fission-fusion; and multi-male, single-female. These types are discussed below.<\/p>\n<h4 class=\"import-Normal\"><em>Solitary<\/em><\/h4>\n<p class=\"import-Normal\">Recall that the term <em>solitary<\/em> is used to describe species in which individuals do not live or travel together with other members of the same species, except for mothers and unweaned offspring. Males typically occupy a large home range or territory that overlaps the home ranges of multiple females, with whom they mate (Figure 7.10a). Because one male mates with multiple females, the mating system of solitary primates is polygyny. Social interactions between adults are limited but because some males do not get to mate, competition between males is intense. When males compete physically, they benefit from large body size and weaponry. The result is<strong> <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1046\">sexual dimorphism<\/a><\/strong>, when males and females look different from one another. Both males and females disperse, although males move farther from their mother than females. The nocturnal West African potto (<em>Perodicticus <\/em><em>potto<\/em>; Figure 7.10b) is solitary. Bornean orangutans, which are diurnal, are also solitary.<\/p>\n<\/div>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_210-9\" aria-describedby=\"caption-attachment-210-9\" style=\"width: 620px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-197\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.10-01-1.jpg\" alt=\"Grouping pattern description is available in caption. A potto in a tree at night also shown.\" width=\"620\" height=\"272\" \/><figcaption id=\"caption-attachment-210-9\" class=\"wp-caption-text\">Figure 7.10a-b: Illustration of a solitary species\u2019 grouping pattern is shown on the left. Key: square = adult male; dot = adult female; open circle represents the outline of the male\u2019s home range; open oval represents individual female home ranges. The West African potto is a solitary primate (right). <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. Polygyny in a Solitary Social System original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Enstam Jaffe is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>. b. <a href=\"https:\/\/www.flickr.com\/photos\/nikborrow\/31307385633\">West African Potto Perodicticus potto Kakum National Park, Ghana<\/a> by<a href=\"https:\/\/www.flickr.com\/people\/nikborrow\/\"> Nik Barrow<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/2.0\/\"> CC BY-NC 2.0 License<\/a>.<\/figcaption><\/figure>\n<div class=\"__UNKNOWN__\">\n<h4 class=\"import-Normal\"><em>Single-Male, Single-Female<\/em><\/h4>\n<p class=\"import-Normal\">Primate species in which an adult male and adult female live together with their dependent offspring have a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1048\">single-male, single-female<\/a><\/strong> social system, sometimes referred to as a \u201cfamily,\u201d with group sizes between two and five individuals. The adult male and adult female engage in behaviours that strengthen their social relationship, or <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1050\">pair bond<\/a><\/strong>, including mutual grooming and resting together. The pair defend a territory (Figure 7.11a) and keep same-sex individuals away from their mate. The adult male and adult female mate with each other, so the mating system is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1052\">monogamy<\/a><\/strong>, although mating outside the pair bond may occur. Species with monogamous mating systems are usually <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1054\">sexually monomorphic<\/a><\/strong> (males and females look similar) because competition for mates is relaxed since most males are able to obtain a mate. Males are usually confident that they are the father of their mate\u2019s infant, so they help with offspring care by carrying the infant when it is not nursing. Once offspring are sexually mature, both males and females disperse. As with solitary species, males disperse farther from their parents than females. Bolivian Gray titi monkeys (<em>Plecturocebus donacophilus<\/em>) are an example of a species that has a single-male, single-female social system. One of their signature behaviours is tail twining, when two individuals sit with their tails wrapped around each other (Figure 7.11b). This behaviour reinforces the social bond among family members and is especially common between the adult male and female. Gibbons (<em>Hylobates<\/em>) and owl monkeys (<em>Aotus<\/em>) also live in single-male, single-female groups.<\/p>\n<figure id=\"attachment_198-2\" aria-describedby=\"caption-attachment-198-2\" style=\"width: 608px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-198\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.11-01.jpg\" alt=\"Left: Circle contains one dot (female) and one square (male). Right: Two titi monkeys.\" width=\"608\" height=\"357\" \/><figcaption id=\"caption-attachment-198-2\" class=\"wp-caption-text\">Figure 7.11a-b: Illustration of a single-male, single-female grouping pattern is shown on the left. Key: square = adult male; dot = adult female; open circle represents the outline of the group\u2019s territory, which the bonded pair defend against conspecifics. The titi monkey (right) is an example of a primate species with a single-male, single-female social system. Credit: a. Single-Male, Single-Female Social System original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Enstam Jaffe is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Callicebus-brunneus-London-Zoo.jpg\">Two Red Titi Monkeys (Callicebus cupreus) sitting together with their tails intertwined at the London Zoo<\/a> by<a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Stevenj\"> Steven G. Johnson<\/a> has been modified (cropped) and 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<\/div>\n<div class=\"__UNKNOWN__\">\n<h4 class=\"import-Normal\"><em>Single-Male, Multi-Female<\/em><\/h4>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1056\">Single-male, multi-female<\/a><\/strong> groups consist of one adult male living with multiple adult females and their dependent offspring (Figure 7.12a ) . These groups can range from as few as five or ten individuals to as many as 50. Female social relationships are governed by the female dominance hierarchy. Females are usually philopatric and males disperse. Males who are unable to join a group of females may join a bachelor group with other males. Because a single male mates with multiple females, the mating system is polygyny. Species that form single-male, multi-female groups may or may not defend a territory, but the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1058\">resident male<\/a><\/strong>, who lives with a group of females, is aggressive toward other males, who may try to take over the group and become the new resident male. Competition between males to be the resident male of a group is intense, and these species usually display sexual dimorphism, with males being larger than females and possessing large canines. Hanuman langurs (<em>Semnopithecus entellus<\/em>) of India form single-male, multi-female groups (Figure 7.12b). When a new male takes over a group of females and ousts the former resident male, he may commit <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1060\">infanticide<\/a>, <\/strong>or kill the unweaned infants. This is especially likely if the new resident male has not yet mated with any of the females and thus cannot be the infants\u2019 father. This causes the females, who were nursing, to become sexually receptive sooner, increasing the new resident male\u2019s chances of producing offspring (Sharma, Ram, and \u200b\u200bRaipurohit 2010). Gorillas, patas monkeys, and golden snub-nosed monkeys (<em>Rhinopithecus roxellana<\/em>) also live in single-male, multi-female groups.<\/p>\n<p>&nbsp;<\/p>\n<\/div>\n<figure id=\"attachment_210-10\" aria-describedby=\"caption-attachment-210-10\" style=\"width: 749px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-199\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.12.jpg\" alt=\"Left: Circle contains nine dots and one square; outside are three squares. Right: Adult langur and infant.\" width=\"749\" height=\"295\" \/><figcaption id=\"caption-attachment-210-10\" class=\"wp-caption-text\">Figure 7.12a-b: An illustration of the one-male, multi-female grouping pattern is shown on the left. Key: square = adult male; dot = adult female; open circle represents the outline of the group\u2019s home range (or territory). The Hanuman langur (right) is an example of a species with a one-male, multi-female social system. Credit: a. Single-Male, Multi-Female Social System original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Enstam Jaffe is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>. b. <a href=\"https:\/\/www.pexels.com\/photo\/close-up-photo-of-two-gray-langurs-8642891\/\">Close-up of Two Grey Langurs<\/a> by<a href=\"https:\/\/www.pexels.com\/@amitrai10\/\"> Amit Rai<\/a> has been modified (cropped) and is<a href=\"https:\/\/www.pexels.com\/license\/\"> free to use via Pexels<\/a>.<\/figcaption><\/figure>\n<div class=\"__UNKNOWN__\">\n<h4 class=\"import-Normal\"><em>Multi-Male, Multi-Female<\/em><\/h4>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1062\">Multi-male, multi-female<\/a><\/strong> groups consist of multiple adult males living with multiple adult females and their dependent offspring. Although there is more than one adult male, there are more adult females than adult males in the group (Figure 7.13a). Multi-male, multi-female groups can range in size from about ten to as many as 500 individuals. They occupy a home range but may or may not defend a territory. In groups that contain multiple males and multiple females, it is not possible for one male to monopolize all the matings, so the mating system is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1064\">polygamy<\/a><\/strong>, in which multiple males mate with multiple females. However, this does not mean that all males have an equal opportunity to mate with all females. In multi-male, multi-female groups, both males and females form a dominance hierarchy. The male dominance hierarchy determines their access to females for mating in much the same way that a female dominance hierarchy determines a female\u2019s access to food. Because their place in the hierarchy can affect their reproductive success, males compete with each other, but because it is rare for males to be excluded from mating altogether, the level of competition and degree of sexual dimorphism are less extreme than what we see in polygynous species. Usually, females are philopatric and males disperse. Vervet monkeys (Figure 7.13b), ring-tailed lemurs (<em>Lemur catta<\/em>), white-faced capuchins (<em>Cebus capucinus<\/em>), and black-capped squirrel monkeys (<em>Saimiri boliviensis<\/em>) live in multi-male, multi-female groups.<\/p>\n<\/div>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_210-11\" aria-describedby=\"caption-attachment-210-11\" style=\"width: 585px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-200\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.13.jpg\" alt=\"Circle contains twelve dots and three squares. Right: Two vervet monkeys.\" width=\"585\" height=\"267\" \/><figcaption id=\"caption-attachment-210-11\" class=\"wp-caption-text\">Figure 7.13a-b: An illustration of the multi-male, multi-female grouping pattern is shown on the left. Key: square = adult male; dot = adult female; open circle represents the outline of the group\u2019s home range (or territory). Vervet monkeys (right) are an example of a species that lives in multi-male, multi-female groups. Credit: a. Multi-Male, Multi-Female Social System original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Enstam Jaffe is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>. b. <a href=\"https:\/\/www.flickr.com\/photos\/berniedup\/6011902081\/\">Vervet Monkeys (Chlorocebus pygerythrus)<\/a> by<a href=\"https:\/\/www.flickr.com\/people\/65695019@N07\"> Bernard DUPONT<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/deed.en\"> CC BY-SA 2.0 License<\/a>.<\/figcaption><\/figure>\n<div class=\"__UNKNOWN__\">\n<h4 class=\"import-Normal\"><em>Fission-Fusion<\/em><\/h4>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1066\">Fission-fusion<\/a><\/strong> is a fluid social system in which the size and composition of the social group changes, with groups splitting (fission) or merging (fusion) depending on food availability (Pinacho-Guendulain &amp; Ramos-Fern\u00e1ndez 2017). When key resources are scarce, individuals spread out (fission) and move and feed individually or in small subgroups (Figure 7.14a). When key food resources are plentiful, individuals come together (fusion) and individuals travel and feed as a more cohesive group (Figure 7.14a). Fission-fusion social structure is believed to reduce feeding competition when resources are scarce. Because group composition changes over time, species with fission-fusion social systems are referred to as a community. Communities consist of multiple adult males, multiple adult females, and offspring, and group size varies but typically ranges from ten to a few dozen individuals. Females typically disperse and males are philopatric. Thus, community males are related and display unusual forms of cooperation. The mating system associated with fission-fusion is polygamy. Because males are not excluded from mating, competition for mates is relaxed and sexual dimorphism is moderate (males are slightly larger than females). Geoffroy\u2019s spider monkeys (<em>Ateles geoffroyi<\/em>) (Figure 7.14b) and chimpanzees both have fission-fusion social system.<\/p>\n<\/div>\n<figure id=\"attachment_210-12\" aria-describedby=\"caption-attachment-210-12\" style=\"width: 720px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-201\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.14.jpg\" alt=\"Diagrams show fission and fusion. Three spider monkeys.\" width=\"720\" height=\"289\" \/><figcaption id=\"caption-attachment-210-12\" class=\"wp-caption-text\">Figure 7.14a-b: An illustration of the fission-fusion grouping pattern appears on the left. The left illustration represents fission, when females travel and feed independently in individual home ranges within the community boundary. The right illustration represents fusion, when community members form a cohesive group. <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>. Key: square = adult male; dot = adult female; open circle represents the outline of the community boundary. Open ovals represent individual female home ranges when the group fissions. Credits: a. Fission-Fusion Social System original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Enstam Jaffe is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>. b. <a href=\"https:\/\/www.flickr.com\/photos\/berniedup\/49562895393\">Geoffry\u2019s Spider Monkeys (Ateles geoffroyi)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/65695019@N07\">Bernard DUPONT<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/deed.en\"> CC BY-SA 2.0 License<\/a>.<\/figcaption><\/figure>\n<div class=\"__UNKNOWN__\">\n<h4 class=\"import-Normal\"><em>Multi-Male, Single-Female<\/em><\/h4>\n<p class=\"import-Normal\">In <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1068\">multi-male, single-female<\/a> <\/strong>groups, two or more males live with one breeding female, her dependent offspring, and non-breeding females (Figure 7.15a). This type of social system is found in the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1070\">callitrichids<\/a><\/strong>, the primate family that includes marmosets (<em>Callithrix<\/em>; Figure 7.15b) and tamarins (<em>Saguinus<\/em>) of Central and South America. Their groups rarely exceed 15 individuals, and each group actively defends their territory from conspecifics. Although more than one adult female may live in the group, the mating system is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1072\">polyandry<\/a><\/strong> because there is only one breeding female who mates with all of the adult males. This is achieved through <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1074\">reproductive suppression<\/a><\/strong>, which involves the breeding female preventing other females from reproducing through physiological and\/or behavioural means (Digby, Ferrari, &amp; Salzman 2011). This limits the opportunities for other females in the group to become pregnant. Instead, these females, and the males in the group, help raise the breeding female\u2019s offspring. This is referred to as <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1076\">cooperative breeding<\/a><\/strong> and usually takes the form of carrying infants, grooming them, and protecting them from danger (de Oliveira Terceiro &amp; Burkart 2019). Because reproductive opportunities for female tamarins and marmosets are limited, they are very competitive, and females are slightly larger than males, which helps them compete for the breeding spot in a group.<\/p>\n<p>&nbsp;<\/p>\n<\/div>\n<figure id=\"attachment_210-13\" aria-describedby=\"caption-attachment-210-13\" style=\"width: 578px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-202\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.15.jpg\" alt=\"Circle contains two squares, two unmarked dots, and a B dot. Right: Marmosets with twins.\" width=\"578\" height=\"269\" \/><figcaption id=\"caption-attachment-210-13\" class=\"wp-caption-text\">Figure 7.15a-b: An illustration of multi-male, single-female grouping pattern appears on the left. Key: square = adult male; B dot = breeding female; unmarked dot = non-breeding female; open circle represents the outline of the group\u2019s territory, which is defended against conspecifics. The common marmoset (Callithrix jacchus) is an example of a primate species that has this type of social system (right). Credit: a. Multi-Male, Single-Female Social System original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Enstam Jaffe is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Family_of_Common_Marmoset_-_REGUA_-_Brazil_MG_9480_(12930855765).jpg\">Family of Common Marmoset - REGUA - Brazil MG 9480 (12930855765)<\/a> by<a href=\"https:\/\/www.flickr.com\/people\/30818542@N04\"> Francesco Veronesi<\/a> has been modified (cropped) 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<div class=\"__UNKNOWN__\">\n<h2 class=\"import-Normal\">Reproductive Strategies<\/h2>\n<p class=\"import-Normal\">Reproductive strategies have evolved to maximize individual reproductive success. These strategies can be divided into those that deal with offspring production and care (parental investment) and those that maximize mating success (sexual selection). Because the reproductive physiology of male and female primates differs, males and females differ with regard to parental investment and sexual selection strategies. Female strategies focus on obtaining the food necessary to sustain a pregnancy and choosing the best male(s) to father offspring. Male strategies focus on gaining access to receptive females.<\/p>\n<h3 class=\"import-Normal\"><strong>Parental Investment<\/strong><\/h3>\n<figure style=\"width: 362px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image9.jpg\" alt=\"Monkey holds baby.\" width=\"362\" height=\"241\" \/><figcaption class=\"wp-caption-text\">Figure 7.16: A female Japanese macaque nursing her infant. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Snow_monkey_baby_milk_time.jpg\">Snow monkey baby milk time<\/a> by Daisuke Tashiro 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>Biologically speaking, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1078\">parental investment<\/a><\/strong> is any time or energy a parent devotes to the current offspring that enhances its survival (and eventual reproductive success) at the expense of the parent\u2019s ability to invest in the next offspring (Trivers 1972). Female primates invest more heavily in offspring than males. Even before conception, females produce energy-containing eggs, and they will be responsible for sustaining a fertilized egg until it implants in the uterus. After that, they invest in pregnancy and lactation (Figure 7.16). Because all of this investment requires a lot of energy, female primates can only produce one offspring (or litter) at a time. A species\u2019 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1080\">interbirth interval<\/a><\/strong>, or the typical length of time between one birth and the next, is determined by the length of time necessary to maximize each offspring\u2019s survival without jeopardizing the female\u2019s ability to produce the greatest number of offspring possible. If a female invests too little (i.e., weans an offspring too early), she may give birth to many offspring, but very few (if any) of them will survive. If she invests too much (i.e., nurses an offspring even after it could be weaned), she ensures the survival of that individual offspring but will not be able to produce very many during her lifetime. To maximize her reproductive success, a female must invest <em>just<\/em> long enough to ensure the greatest number of offspring survive to reproduce. We often think of maternal care as an <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1082\">innate<\/a><\/strong> (or natural), instinctive behaviour. Yet this is not the case. The \u201cSpecial Topic: Is Maternal Behaviour Innate?\u201d dispels the myth that maternal behaviour is solely instinctual and explains how female primates learn to be good mothers.<\/p>\n<h3 class=\"import-Normal\"><strong>Sexual Selection<\/strong><\/h3>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1084\">Sexual selection<\/a><\/strong>, or selection for traits that maximize mating success, comes in two forms. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1086\">Intrasexual selection<\/a><\/strong> is selection for traits that enhance the ability of members of one sex to compete amongst themselves (\u201c<em>intra<\/em>sexual\u201d = within one sex). <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1088\">Intersexual selection<\/a><\/strong> is selection for traits that enhance the ability of one sex to attract the other (\u201c<em>inter<\/em>sexual\u201d = between the sexes).<\/p>\n<p class=\"import-Normal\">Intrasexual selection most often operates on males. In the wild, adult females are either pregnant or lactating for most of their adult lives. So, in a given population, there are usually more males available and willing to mate than there are females. The result? Females are a scarce resource over which males compete. Intrasexual selection favours traits that help a male win fights with other males. In primates, these traits include large body size (Figure 7.17a) and large canines (Figure 7.17b). Because females don\u2019t possess these same traits, males and females of some species look different; that is, they are sexually dimorphic (Figure 7.17a).<\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_207-4\" aria-describedby=\"caption-attachment-207-4\" style=\"width: 691px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-204\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.17.jpg\" alt=\"Male baboon with two females. Adult male baboon.\" width=\"691\" height=\"293\" \/><figcaption id=\"caption-attachment-207-4\" class=\"wp-caption-text\">Figure 7.17a-b: a. Hamadryas baboons (Papio hamadryas) are sexually dimorphic. The male (left) is much bigger than the female (centre) and also has different coloured fur. b. Adult males, like this gelada baboon, also have larger canines than females. Credit: a. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Dierenpark_Emmen_baboon_(2679944324).jpg\">Dierenpark Emmen baboon (2679944324)<\/a> by<a href=\"https:\/\/www.flickr.com\/people\/80538772@N00\"> robin bos<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\"> CC BY 2.0 License.<\/a> b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Olive_Baboon_Papio_anubis_in_Tanzania_3066_Nevit.jpg\">Olive Baboon Papio anubis, Picture Taken in Tanzania<\/a> by Nevit Dilmen has been modified (cropped) and 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\">Intersexual selection also tends to operate on males, selecting traits that make a male more attractive to females. Females, in turn, choose among potential fathers. Because female primates invest more in offspring production and care than males (see the \u201cParental Investment\u201d section, above), it is more costly for them if the offspring dies before maturity or reaches maturity but does not reproduce. Thus, it benefits a female primate to be choosy and try to pick the healthiest male as a mate. Males must display traits that tell a female why she should choose <em>him<\/em>, and not another male, as her mate.<\/p>\n<p class=\"import-Normal\">What traits are female primates looking for? In humans, women may look for a mate who can provide important resources, such as food, paternal care, or protection. This is rare in other primates, though, since most females do not need males to provide resources. More commonly, female primates obtain genetic benefits for their offspring from choosing one male over another. Often the specific criteria by which females select mates is unknown. However, if a female chooses a healthy (as indicated by traits like a plush coat, bright colouration, or large body size) or older male, she may obtain genes for her offspring that code for health or long life. If a male\u2019s rank is determined by competitive ability that has a genetic component, females who choose males who win fights may acquire these genes (and qualities) for their offspring. Females in some species appear to prefer new immigrants, sometimes even \u201csneaking\u201d copulations with males who are not established members of their groups. Such a preference may provide their offspring with novel genes and increase genetic variation (for more about the importance of genetic variation, see Chapter 4). Female choice is often more subtle than male-male competition, so it can be more difficult to study. However, as more research is conducted, we continue to improve our understanding of the ways that female primates exert their choice.<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Is Maternal Behaviour Innate?<\/h2>\n<p>Zoos almost always have nurseries where infants are cared for by zookeepers if their mothers will not care for them (Figure 7.18). These exhibits are among the most popular because the babies are so cute and so much fun to watch. And the caretaking positions in zoo nurseries are often among the most coveted by zoo personnel for the same reasons. But if maternal behaviour is instinctive, why do zoo nurseries even exist? The answer is that in many species, including primates, maternal behaviour is not purely instinctual; it is dependent on <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1090\">social learning<\/a><\/strong> (behaviour learned by observing and imitating others), as well.<\/p>\n<figure style=\"width: 433px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image13-1.png\" alt=\"Newborn orangutan feeding from a bottle.\" width=\"433\" height=\"336\" \/><figcaption class=\"wp-caption-text\">Figure 7.18: Newborn orangutan at Audubon Zoo being bottle-fed. Credit: <a href=\"https:\/\/newsroom.audubonnatureinstitute.org\/critically-endangered-orangutan-gives-birth-at-audubon-zoo\/\">Newborn orangutan born at Audubon Zoo being bottle fed<\/a> (2022) by<a href=\"https:\/\/audubonnatureinstitute.org\/\"> Audubon Nature Institute<\/a> is used by permission.<\/figcaption><\/figure>\n<p>Captive female primates, including gorillas and chimpanzees, who have not had the opportunity to observe their mother or other females care for infants do not know how to care for their own offspring. Although it is preferred that the primate mother care for her own infant, there are cases when she will not and humans must step in to ensure the offspring survives. When hand-rearing by humans is necessary, the infant is returned to the group as soon as possible in the hopes that it will learn species-typical behaviour from its mother and other conspecifics. Observations such as these indicate that maternal behaviour is learned, not innate, and that maternal care is critically important to the social and psychological development of young primates.<\/p>\n<\/div>\n<h2 class=\"import-Normal\">Communication<\/h2>\n<p class=\"import-Normal\">In its most basic form, communication occurs when one individual (the sender) emits a signal that conveys information, which is detected by another individual (the receiver). We have discussed several aspects of primate sociality in this chapter, all of which require the communication of information between individuals. But exactly <em>how<\/em> does a female chimpanzee communicate her sexual availability? <em>How<\/em> does a vervet monkey communicate the approach of a leopard or that a python is nearby? <em>How<\/em> do solitary, nocturnal primates, like the slow loris, communicate information about themselves to conspecifics? Primate communication comes in four forms: vocal, visual, olfactory, and tactile. Species vary in their reliance on each.<\/p>\n<h3 class=\"import-Normal\"><strong>Vocal Communication<\/strong><\/h3>\n<p class=\"import-Normal\">Primates use sound to communicate danger or threats, to claim and maintain a territory, or make contact with other group members. Alarm calls are given in response to predators. In some cases, alarm calls are used to alert members of the group to the presence of a predator so they can take evasive action. In other cases, they are directed at the predator itself, signaling that it has been detected. You can learn more about alarm calls as forms of vocal communication in the highlight box in this chapter entitled \u201cDig Deeper: Alarm Calls: Signals to Friends or Foes?.\u201d<\/p>\n<p class=\"import-Normal\">Loud calls are designed to travel great distances and are used in territorial defence by many primate species including indris (<em>Indri indri<\/em>), orangutans, gibbons, and howler monkeys (<em>Alouatta<\/em>). In dense forest, where visual communication can be difficult, loud calls can be useful in signaling to conspecifics that a group or individual occupies a specific area. Howler monkeys are named for their loud calls, or \u201croars,\u201d which can be heard one kilometre or more away (Sch\u00f6n Ybarra 1986). Howler monkey roars may act to maintain distance between neighbouring groups or keep extragroup males from entering the home range (1986).<\/p>\n<p class=\"import-Normal\">Other vocalizations are intended to communicate with individuals in one\u2019s own group. These include vocalizations given as part of threat displays or dominance interactions, as well as contact calls that provide information about one\u2019s location to other group members. Chacma baboons (<em>Papio ursinus<\/em>) have a rich repertoire of vocalizations for communicating with other group members (Fischer Et al. 2008). Adult males give specific vocalizations during threat displays and physical confrontations. Subordinates \u201cscreech\u201d when retreating from a dominant individual, signaling submission. Since baboons rely on membership in their group for finding food and detecting predators, a baboon separated from his group will vocalize in an attempt to regain contact. Young baboons emit their own contact calls when separated from their mothers.<\/p>\n<h3 class=\"import-Normal\"><strong>Visual Communication<\/strong><\/h3>\n<figure id=\"attachment_207-5\" aria-describedby=\"caption-attachment-207-5\" style=\"width: 493px\" class=\"wp-caption alignleft\"><img class=\"wp-image-206\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.19.jpg\" alt=\"Female baboon with sexual swelling. Male and female baboon.\" width=\"493\" height=\"209\" \/><figcaption id=\"caption-attachment-207-5\" class=\"wp-caption-text\">Figure 7.19a-b: Two female hamadryas baboons. The female on the left has a sexual swelling while the female on the right (in foreground, with infant clinging to her belly) does not. An adult male is behind her. Credit: a. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Sexual_swelling_in_female_Hamadryas_baboon.jpg\">Sexual swelling in female Hamadryas baboon<\/a> by<a href=\"https:\/\/www.flickr.com\/photos\/mamoritai\/\"> Mamoritai<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/legalcode\"> CC BY-SA 2.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Hamadryas_baboon_at_Giza_Zoo_by_Hatem_Moushir_36.JPG\">Hamadryas baboon at Giza Zoo by Hatem Moushir 36<\/a> by<a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Hatem_Moushir\"> Hatem Moushir<\/a> has been modified (cropped) and 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><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1092\">Visual communication<\/a><\/strong>, which involves signals that can be seen, is an important component of nonhuman primate behaviour, alone or in combination with other forms of communication. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1094\">Piloerection<\/a><\/strong>, or raising one\u2019s hair or fur, is used in aggressive interactions to make an individual appear larger than it actually is. Female macaques (<em>Macaca<\/em>), baboons (<em>Papio<\/em>), and chimpanzees, signal sexual receptivity through changes in the size, shape, and, often, colour of their hindquarters, called a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1096\">sexual swelling<\/a><\/strong> (Figure 7.19a). The sexual swelling reaches its maximum size at ovulation. When females are not receptive, either because they are pregnant or are nursing, they do not display a sexual swelling (Figure 7.19b). Thus, the presence or absence of a sexual swelling signals a female\u2019s reproductive state.<\/p>\n<p>Monkeys and apes use diverse facial expressions in visual communication. Showing your teeth in a \u201csmile\u201d sends a signal of friendship in humans. Displaying teeth in this way is a sign of anxiety or fear in primates. That male mandrill you see \u201cyawning\u201d at your local zoo is actually displaying his teeth to signal tension or to threaten a rival (Figure 7.20a). In addition to showing their canines, male gelada baboons use \u201clip flips,\u201d in which the gums and teeth are exposed by flipping the upper lip up over the nostrils (Figure 7.20b), and \u201craised eyelids,\u201d in which the pale eyelids are exposed by pulling the scalp back as threatening gestures (Aich, Moos-Heilen, &amp; Zimmerman 1990). Submissive males respond by fleeing or presenting their hindquarters.<\/p>\n<figure id=\"attachment_207-6\" aria-describedby=\"caption-attachment-207-6\" style=\"width: 597px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-207\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.20-1.jpg\" alt=\"Adult male mandrill and adult male hamadryas baboon yawning.\" width=\"597\" height=\"310\" \/><figcaption id=\"caption-attachment-207-6\" class=\"wp-caption-text\">Figure 7.20a-b: Males use visual displays to communicate with other males. The male mandrill (left) is yawning to display his canines, and the male gelada baboon (right) enhances the yawn by flipping his upper lip back and raising his eyelids. Credit: a. <a href=\"https:\/\/pxhere.com\/en\/photo\/559944\">Mandrill<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/mathiasappel\/with\/24639723420\/\">Mathias Appel<\/a> has been modified (cropped) and designated to the <a href=\"https:\/\/creativecommons.org\/share-your-work\/public-domain\/cc0\/\">public domain (CC0)<\/a>. b.<a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:BabouinGeladaAuReveil.JPG\"> BabouinGeladaAuReveil<\/a> by<a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:BluesyPete\"> BluesyPete<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\"> CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<figure id=\"attachment_209-2\" aria-describedby=\"caption-attachment-209-2\" style=\"width: 414px\" class=\"wp-caption alignleft\"><img class=\"wp-image-208\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.22.jpg\" alt=\"A bald uakari. A spider monkey.\" width=\"414\" height=\"154\" \/><figcaption id=\"caption-attachment-209-2\" class=\"wp-caption-text\">Figure 7.22a-b: Many monkey species have colourful faces, including the bald uakari (Cacajao calvus; left) and the white-bellied spider monkey (Ateles belzebuth) (right). Credit: a. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Uakari.jpg\">Uakari<\/a> by Coada dragos has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\"> CC BY-SA 4.0 License<\/a>. 6.22b <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Ateles_belzebuth_(White-bellied_spider_monkey)_2.jpg\">Ateles belzebuth (White-bellied spider monkey) 2<\/a> by<a href=\"https:\/\/www.flickr.com\/photos\/ewas-world\/\"> Ewa<\/a> (username: Ewcek65) has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\"> CC BY 2.0 License<\/a>.<\/figcaption><\/figure>\n<figure style=\"width: 163px\" class=\"wp-caption alignright\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-3.png\" alt=\"A male mandrill\u2019s face.\" width=\"163\" height=\"246\" \/><figcaption class=\"wp-caption-text\">Figure 7.21: The colourful face of the male mandrill provides information about health and fitness to other mandrills.Credit: <a href=\"https:\/\/www.flickr.com\/photos\/mathiasappel\/24842082402\/\">Mandrill<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/mathiasappel\/with\/24639723420\/\">Mathias Appel<\/a> has been modified (cropped) and designated to the <a href=\"https:\/\/creativecommons.org\/share-your-work\/public-domain\/cc0\/\">public domain (CC0)<\/a>.<\/figcaption><\/figure>\n<p>Primates also communicate through colour. In female and male mandrills, facial colouration provides information about an individual\u2019s health, competitive ability, and reproductive state to conspecifics (Figure 7.21; Setchell &amp;t al. 2008; Setchell, Wickings, &amp; Knapp 2006). Variation in facial colouration among monkeys of Central and South America ranges from very simple (Figure 7.22a) to complex (Figure 7.22b). Species living with larger numbers of other primate species have evolved more complex facial colouration patterns, suggesting that this trait evolved as a form of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1098\">species recognition<\/a><\/strong>, or the ability to differentiate conspecifics from members of other species (Santana, Lynch Alfaro, &amp; Alfaro 2012).<\/p>\n<h3 class=\"import-Normal\"><strong>Olfactory Communication<\/strong><\/h3>\n<p class=\"import-Normal\">All primates use scent to communicate. Females secrete chemicals from their <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1100\">anogenital<\/a> <\/strong>region (the area of the anus and genitals) that provide males with information about their reproductive state. In some species, like macaques and chimpanzees, this olfactory signal is enhanced by a sexual swelling, as discussed above. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1102\">Olfactory communication<\/a><\/strong>, or communicating through scent, is particularly important for monkeys of Central and South America, lemurs, and lorises. Male and female common squirrel monkeys (<em>Saimiri sciureus<\/em>) (Figure 7.23a) engage in \u201curine washing,\u201d in which an individual urinates on its hands and feet and then uses them to spread urine all over its body. Urine washing may be used to mark trails for others to follow, to control body temperature, as part of dominance displays, or to communicate reproductive state (Boinski 1992). During aggressive interactions with other males, male ring-tailed lemurs rub their tails with scent from glands on their wrists and chests. They use their \u201cperfumed\u201d tails in aggressive interactions with other males, who may respond by waiving their own scented tail, with physical aggression, or by fleeing (Jolly 1966). Males also waive their tails, saturated in scent, to attract females (Shirasu et al. 2020). Males use scent glands in their wrists to mark territorial boundaries (Figure 7.23b; Mertl-Millhollen 1988).<\/p>\n<\/div>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_210-14\" aria-describedby=\"caption-attachment-210-14\" style=\"width: 641px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-210\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.23.jpg\" alt=\"A squirrel monkey. A ring-tailed lemur.\" width=\"641\" height=\"332\" \/><figcaption id=\"caption-attachment-210-14\" class=\"wp-caption-text\">Figure 7.23a-b: Some primates, like the common squirrel monkey (left) and the ring-tailed lemur (right), communicate using scent. Credit: a. <a href=\"https:\/\/www.flickr.com\/photos\/rubund\/6337874822\/\">Saimiri sciureus<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/rubund\/\">Ruben Undheim<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/\">CC BY-SA 2.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Lemur_catta_004.jpg\">Lemur catta 004<\/a> by <a href=\"https:\/\/en.wikipedia.org\/wiki\/User:Maky\">Maky<\/a> has been modified (cropped) and 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<div class=\"__UNKNOWN__\">\n<h3 class=\"import-Normal\"><strong>Tactile Communication<\/strong><\/h3>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1104\">Tactile communication<\/a><\/strong>, or communicating through touch, is very important in all primate species. Physical contact is used to comfort and reassure, is part of courtship and mating, and is used to establish dominance and alliances. Grooming is an important and clearly enjoyable form of tactile communication for all primates (Figure 7.24). Not only does grooming serve to clean the skin and fur, removing parasites and debris, but it is an important affiliative behaviour that helps reinforce social bonds, repair relationships, and cement alliances.<\/p>\n<figure style=\"width: 674px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image1-8.png\" alt=\"Four primate species grooming.\" width=\"674\" height=\"478\" \/><figcaption class=\"wp-caption-text\">Figure 7.24: Examples of grooming in Japanese macaques (upper left), tufted capuchins (Sapajus apella) (upper right), gelada baboons (lower left), and black-and-white ruffed lemurs (Varecia variegata; lower right). Credit: Examples of grooming original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Jaffe 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:Yakushima_macaques_grooming_each_other.jpg\">Yakushima macaques grooming each other<\/a> by<a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Grendelkhan\"> Grendelkhan<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Tufted_capuchin_monkeys_grooming_session_III.jpg\">Tufted capuchin monkeys grooming session III<\/a> by Adrian Soldati, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>;<a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Baboons_Wunania_012018.jpg\"> Baboons Wunania 012018<\/a> by Kim Toogood, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>;<a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Black-and-white_ruffed_lemur_03.jpg\"> Black-and-white ruffed lemur 03<\/a> by Mattis2412, <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/deed.en\">public domain (CC0 1.0)<\/a>].<\/figcaption><\/figure>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Dig Deeper: Alarm Calls: Signals to Friends or Foes?<\/h2>\n<p class=\"import-Normal\">Alarm calls are common among group-living primates. They often serve to notify conspecifics of potential danger, as is the case with vervet monkeys. Research has shown that: (1) vervets classify predators based on hunting style; (2) alarm calls convey information to other vervets about that hunting style; and (3) other vervets respond in ways appropriate for evading that type of predator (Seyfarth, Cheney, &amp; Marler 1980a). When a vervet gives a \u201cleopard\u201d alarm call (directed at mammalian carnivores like leopards, Figure 7.25a), monkeys on the ground climb the nearest tree, while monkeys already in trees stay there or climb higher. Since most mammalian carnivores hunt on the ground, getting into, and staying in, a tree is the best option for escape. When the \u201csnake\u201d alarm call is given, vervets stand on their hind legs and look down at the ground (Figure 7.25b). Since snakes are not pursuit predators, locating them quickly so as to avoid them is the best strategy. Lastly, when an \u201ceagle\u201d alarm call is given, vervets look up or run into bushes, both of which are useful responses for avoiding hawks and eagles, which attack from above (Figure 7.25c). Vervets clearly understand the meaning of each type of alarm call, as they respond appropriately even when they do not see the actual predator (Seyfarth, Cheney, &amp; Marler 1980b). Such <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1106\">semantic communication<\/a><\/strong>, which involves the systematic use of signals to refer to objects in the environment, was once believed to be unique to humans. It may be a precursor to the symbolic capacities of human language.<\/p>\n<figure style=\"width: 482px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5-1.jpg\" alt=\"Primate in a tree views a leopard.\" width=\"482\" height=\"344\" \/><figcaption class=\"wp-caption-text\">Figure 7.25a<\/figcaption><\/figure>\n<figure style=\"width: 484px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image2-2.jpg\" alt=\"Primate views snake on the ground.\" width=\"484\" height=\"338\" \/><figcaption class=\"wp-caption-text\">Figure 7.25b<\/figcaption><\/figure>\n<figure style=\"width: 481px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-2.jpg\" alt=\"Primate on the ground sees bird.\" width=\"481\" height=\"517\" \/><figcaption class=\"wp-caption-text\">Figure 7.25c\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 Figure 7.25a-c: Vervet monkeys respond in different ways to alarm calls for each of their three main predators (leopards, snakes, and eagles) which are appropriate to predator hunting strategies. Credit: Vervet Monkey Alarm Calls by Mary Nelson, original to Explorations: An Open Invitation to Biological Anthropology, 2nd edition, 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\">Research on other African monkeys indicates that some species use alarm calls to signal to the predator that it has been detected. Diana monkeys (<em>Cercopithecus diana<\/em>) give alarm calls to leopards (<em>Panthera pardus<\/em>) but not chimpanzees (Zuberb\u00fchler, No\u00eb, &amp; Seyfarth 1997). Because leopards are stealth predators, they rely on the element of surprise to sneak up on their prey (Figure 7.26a). Alarm calling at leopards appears to tell the leopard that it has been seen and therefore its chance of success will be low. Research shows leopards are more likely to stop hunting after an alarm call has been emitted. Unlike leopards, chimpanzees are pursuit predators and may even use alarm calls to locate potential prey (Figure 7.26b). With such a predator, prey are better off remaining as silent as possible so as not to alert the predator to their location (Zuberb\u00fchler Et al. 1999).<\/p>\n<figure id=\"attachment_215-2\" aria-describedby=\"caption-attachment-215-2\" style=\"width: 1907px\" class=\"wp-caption alignnone\"><img class=\"wp-image-215 size-full\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.26.jpg\" alt=\"Leopard crouches in grass. Chimpanzee looks up. \" width=\"1907\" height=\"589\" \/><figcaption id=\"caption-attachment-215-2\" class=\"wp-caption-text\">Figure 7.26a-b: Because leopards (left) and chimpanzees (right) hunt differently, Diana monkeys react differently to them. Credit: a. <a href=\"https:\/\/www.flickr.com\/photos\/thimindu\/5842997328\">Crouching Leopard<\/a> by<a href=\"https:\/\/www.flickr.com\/photos\/thimindu\/\"> Thimindu Goonatillake<\/a> is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/\"> CC BY-SA 2.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Chimpanzee_in_the_wild.jpg\">Chimpanzee in the wild<\/a> by D.G. Kulakov 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\">\n<\/div>\n<h2 class=\"import-Normal\"><span style=\"text-align: initial;font-size: 1em\">The Question of Culture<\/span><\/h2>\n<p class=\"import-Normal\">It may be surprising in a chapter on nonhuman primates to see a discussion of culture. After all, culture is considered by many, including cultural anthropologists, to be a distinguishing characteristic of humans. Indeed, some anthropologists question claims of culture in primates and other animals. Definitions of animal culture focus on specific behaviours that are unique to one population. Anthropological definitions of human culture emphasize shared ideology (e.g., values, morals, beliefs) and symbols, not just behaviour. Using this definition, some cultural anthropologists view primates as lacking culture because of the absence of symbolic life (e.g., religion). However, the longer we study primate groups and populations, the more insight we gain into primate behavioural variation. If we define <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1108\">culture<\/a><\/strong> as the transmission of behaviour from one generation to the next through social learning, then we must view at least some of the behavioural variation we see in primates as forms of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1110\">cultural tradition<\/a><\/strong>, or a distinctive pattern of behaviour shared by multiple individuals in a social group that persists over time (Whiten 2001).<\/p>\n<h3 class=\"import-Normal\"><strong>Chimpanzee Culture<\/strong><\/h3>\n<p class=\"import-Normal\">Due to both their high level of intelligence and the large number of long-term studies on several different populations, chimpanzees provide the best example of cultural tradition in primates. Chimpanzees express cultural variation in multiple behavioural patterns, ranging from population-specific prey preferences and hunting strategies to tool-use techniques and social behaviours. For example, in Tanzania, chimpanzees fish for termites by stripping twigs and then poking the twigs into termite mounds. The termites react to the \u201cinvasion\u201d by attacking the twig. The chimpanzee pulls the twig out, termites attached, and eats them. In Gambia, they use modified twigs to extract honey from holes in trees. In Fongoli, S\u00e9n\u00e9gal, chimpanzees use sticks as \u201cspears\u201d that they stab into tree cavities to hunt for galagos (Figure 7.27). Multiple chimpanzee populations use a \u201chammer and anvil\u201d to crack open nuts, but the specific techniques differ. Because the cultural traditions are so diverse and unique, if a researcher can observe enough of a chimpanzee\u2019s behaviour, it is possible to assign that individual to a specific community, much in the same way a human being can be associated with a specific culture based on his or her behaviour (Whiten 2011).<\/p>\n<figure style=\"width: 800px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image3-4.jpg\" alt=\"Chimpanzees hunting galagos by poking them with a stick. \" width=\"800\" height=\"453\" \/><figcaption class=\"wp-caption-text\">Figure 7.27a-d: Tool-assisted hunting by a chimpanzee at Fongoli, S\u00e9n\u00e9gal. An adult male chimpanzee uses a tree branch with a modified end to (a\u2013c) stab into a cavity within a hollow tree branch that houses a galago. He ultimately captures the galago as (d) his adolescent brother looks on. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Pan_troglodytes,_tool_use_in_Senegal.jpg\">Pan troglodytes, tool use in Senegal<\/a> by<a href=\"https:\/\/royalsocietypublishing.org\/content\/2\/4\/140507\"> J. D. Pruetz, P. Bertolani, K. Boyer Ontl, S. Lindshield, M. Shelley, and E. G. Wessling<\/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\">How do chimpanzee cultures develop, and how does cultural transmission occur? Although we do not know for sure how chimpanzee cultural traditions develop initially, it is possible that different groups invent, either accidentally or deliberately, certain behaviours that other individuals copy. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1112\">Immigration<\/a><\/strong>, or movement of an individual into a new group or community, is an important avenue of cultural transmission in chimpanzees, much as it is between human cultures. Immigrants (typically females) may bring cultural traditions to their new community, which residents observe and learn. Conversely, immigrants may observe and learn a cultural tradition practiced in their new community (Whiten 2011).<\/p>\n<h3 class=\"import-Normal\"><strong>Cultural Transmission in Macaques<\/strong><\/h3>\n<p class=\"import-Normal\">Two monkey species are well-known for behavioural variation that has been called \u201cpre-cultural\u201d by some primatologists: Japanese macaques and tufted capuchins (<em>Sapajus apella<\/em>). The transmission of unique <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1114\">foraging<\/a><\/strong> (the act of searching for food) behaviours through the members of a provisioned group of Japanese macaques on Koshima Island is well known (Matsuzawa 2015). In an effort to keep the monkeys nearby, researchers provided them with piles of sweet potatoes. A juvenile female named Imo spontaneously washed a muddy sweet potato in a stream. This new food-processing technique first spread among other juveniles and then gradually to older individuals. Within 30 years, it had spread across generations, and 46 of 57 monkeys in the group engaged in the behaviour. Another example comes from a group living far to the north, in Shiga-Heights, Nagano Prefecture. Researchers used apples to entice Japanese macaques to the area. Within a few years, monkeys visited the area regularly and were observed playing with the water in the hot springs. Soon, they climbed into the hot springs and learned to immerse themselves to keep warm and reduce stress when not foraging (Figure 7.28; Matsuzawa 2018; Takeshita Et al. 2018; recall also our discussion of hot spring use as an example of analogous traits at the beginning of this chapter). These examples share several characteristics with human culture, including invention or modification of behaviour, transmission of behaviour between individuals, and the persistence of the behaviour across generations (McGrew 1998).<\/p>\n<figure style=\"width: 477px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image6-4.png\" alt=\"Two monkeys in a hot spring.\" width=\"477\" height=\"292\" \/><figcaption class=\"wp-caption-text\">Figure 7.28: Hot spring use by Japanese macaques is a culturally transmitted behaviour. Credit: <a href=\"https:\/\/www.flickr.com\/photos\/peterthoeny\/32160301021\">Oooh, This Feels Sooo Good!<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/peterthoeny\/\">Peter Theony - Quality HD Photography<\/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<h2 class=\"import-Normal\">Summary<\/h2>\n<p class=\"import-Normal\">Primates are socially complex, extremely intelligent, and highly adaptable. In this chapter we discussed aspects of primate ecology, including how body size and characteristics of food affect what primates eat and how primates interact with other species in their environment. We examined why primates live in groups, the types of groups in which they are found, and the reproductive strategies used by males and females to maximize reproductive success. Like other aspects of their behaviour, primate communication is varied and complex, and we discussed how primates communicate using vocal, visual, olfactory, and tactile signals. Finally, we explored the question of culture among nonhuman primates and learned that some species have cultural traditions, distinctive patterns of behaviour shared by multiple individuals in a social group that persist over time. Humans and other primates are similar in many ways. Learning about principles of primate ecology and behaviour can help us better understand our own behaviour and the behaviours of our extinct relatives.<\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Review Questions<\/h2>\n<ul>\n<li class=\"import-Normal\">If anthropology is the study of humans, why do some anthropologists study primates?<\/li>\n<li class=\"import-Normal\">How does a primate\u2019s ecology affect their diet and interactions with other organisms?<\/li>\n<li class=\"import-Normal\">Why do primates live in groups and in what types of groups do they live?<\/li>\n<li class=\"import-Normal\">What is parental investment and sexual selection?<\/li>\n<li class=\"import-Normal\">What are some examples of primate communication?<\/li>\n<li class=\"import-Normal\">What is the evidence for cultural traditions in primates and how do primatologists think cultural transmission occurs in primates?<\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\">Key Terms<\/h2>\n<p class=\"import-Normal\"><strong>Abundance<\/strong>: How much food is available in a given area.<\/p>\n<p class=\"import-Normal\"><strong>Adaptation<\/strong>: A trait with a function.<\/p>\n<p class=\"import-Normal\"><strong>Affiliative<\/strong>: Nonaggressive social interactions and associations between individuals.<\/p>\n<p class=\"import-Normal\"><strong>Agonistic<\/strong>: Conflict; aggressive interactions between individuals.<\/p>\n<p class=\"import-Normal\"><strong>Alarm calling<\/strong>: Vocalizations emitted by social animals in response to danger.<\/p>\n<p class=\"import-Normal\"><strong>Analogy<\/strong>: A similar trait found in different species that arose independently.<\/p>\n<p class=\"import-Normal\"><strong>Anogenital<\/strong>: Relating to the anus and genitals.<\/p>\n<p class=\"import-Normal\"><strong>Breeding season<\/strong>: The time of year when females are receptive to mating.<\/p>\n<p class=\"import-Normal\"><strong>Callitrichids<\/strong>: The primate family that includes marmosets and tamarins.<\/p>\n<p class=\"import-Normal\"><strong>Carnivores<\/strong>: Organisms whose diet consists primarily of animal tissue.<\/p>\n<p class=\"import-Normal\"><strong>Coalition<\/strong>: A temporary alliance between individuals.<\/p>\n<p class=\"import-Normal\"><strong>Community ecology<\/strong>: The branch of ecology that deals with the relationships and interactions between different organisms that occupy the same habitat.<\/p>\n<p class=\"import-Normal\"><strong>Comparison<\/strong>: An examination of the similarities and differences between two things, such as two primate species.<\/p>\n<p class=\"import-Normal\"><strong>Conspecifics<\/strong>: Members of the same species.<\/p>\n<p class=\"import-Normal\"><strong>Cooperative breeding<\/strong>: When individuals other than the mother and father help raise the offspring.<\/p>\n<p class=\"import-Normal\"><strong>Crypsis<\/strong>: The ability to avoid detection by other organisms, such as predators.<\/p>\n<p class=\"import-Normal\"><strong>Cultural tradition<\/strong>: A distinctive pattern of behaviour shared by multiple individuals in a social group, which persists over time and is acquired through social learning.<\/p>\n<p class=\"import-Normal\"><strong>Culture<\/strong>: The transmission of behaviour from one generation to the next through observation and imitation.<\/p>\n<p class=\"import-Normal\"><strong>Decolonize<\/strong>: Understanding and highlighting the theory and research of non-Western individuals and perspectives.<\/p>\n<p class=\"import-Normal\"><strong>Descendant<\/strong>: A species that comes after the ancestor species.<\/p>\n<p class=\"import-Normal\"><strong>Direct competition:<\/strong> Competition that involves physical interaction between individuals, such as fighting.<\/p>\n<p class=\"import-Normal\"><strong>Dispersal<\/strong>: To leave one\u2019s group or area. This may or may not involve joining another group.<\/p>\n<p class=\"import-Normal\"><strong>Distribution<\/strong>: How food is spread out.<\/p>\n<p class=\"import-Normal\"><strong>Diurnal<\/strong>: Active during the day.<\/p>\n<p class=\"import-Normal\"><strong>Dominance hierarchy<\/strong>: The ranked organization of individuals established by the outcome of aggressive-submissive interactions.<\/p>\n<p class=\"import-Normal\"><strong>Dominant<\/strong>: Being of high rank.<\/p>\n<p class=\"import-Normal\"><strong>Ecology<\/strong>: The relationship between organisms and their physical surroundings.<\/p>\n<p class=\"import-Normal\"><strong>Ecotourism<\/strong>: A form of tourism that focuses on nature-based attractions to provide learning opportunities and that uses economically and ecologically sustainable practices.<\/p>\n<p class=\"import-Normal\"><strong>Ethology<\/strong>: The study of animal behaviour.<\/p>\n<p class=\"import-Normal\"><strong>Fission-fusion<\/strong>: Societies in which group composition is flexible, such as chimpanzee and spider monkey societies. Individuals may break up into smaller feeding groups (fission) and combine into larger groups (fusion).<\/p>\n<p class=\"import-Normal\"><strong>Fitness<\/strong>: An individual\u2019s ability to survive and reproduce relative to other members of the same species.<\/p>\n<p class=\"import-Normal\"><strong>Folivores<\/strong>: Organisms whose diet consists primarily of leaves.<\/p>\n<p class=\"import-Normal\"><strong>Foraging<\/strong>: The act of searching for food.<\/p>\n<p class=\"import-Normal\"><strong>Frugivores<\/strong>: Organisms whose diet consists primarily of fruit.<\/p>\n<p class=\"import-Normal\"><strong>Grooming<\/strong>: Picking through the fur of another individual for cleaning or bonding purposes.<\/p>\n<p class=\"import-Normal\"><strong>Heterospecifics<\/strong>: Members of different species.<\/p>\n<p class=\"import-Normal\"><strong>Holism<\/strong>: The idea that the parts of a system interconnect and interact to make up the whole.<\/p>\n<p class=\"import-Normal\"><strong>Home range<\/strong>: The area that a group or individual uses over a given period of time (often over a year).<\/p>\n<p class=\"import-Normal\"><strong>Homology<\/strong>: A similar trait found in different species because it was inherited from a common ancestor.<\/p>\n<p class=\"import-Normal\"><strong>Immigration<\/strong>: Movement of an individual into a new group or community.<\/p>\n<p class=\"import-Normal\"><strong>Indirect competition<\/strong>: Competition that does not involve physical interaction between individuals, such as eating food before another individual arrives at the food site.<\/p>\n<p class=\"import-Normal\"><strong>Infanticide<\/strong>: The killing of infants of one\u2019s own species.<\/p>\n<p class=\"import-Normal\"><strong>Innate<\/strong>: Natural; as in behaviour that comes naturally.<\/p>\n<p class=\"import-Normal\"><strong>Insectivores<\/strong>: Organisms whose diets consist primarily of insects.<\/p>\n<p class=\"import-Normal\"><strong>Interbirth interval<\/strong>: The typical length of time between one birth and the next for a species.<\/p>\n<p class=\"import-Normal\"><strong>Intersexual selection<\/strong>: The selection for traits that enhance the ability of the members of one sex to attract the attention of the other.<\/p>\n<p class=\"import-Normal\"><strong>Intrasexual selection<\/strong>: Selection for traits that enhance the ability of members of one sex to compete amongst themselves.<\/p>\n<p class=\"import-Normal\"><strong>Mating system<\/strong>: A way of describing which male(s) and female(s) mate.<\/p>\n<p class=\"import-Normal\"><strong>Metabolism<\/strong>: The chemical changes that take place in an organism that turn nutrients into energy.<\/p>\n<p class=\"import-Normal\"><strong>Mobbing<\/strong>: Cooperatively attacking or harassing a predator.<\/p>\n<p class=\"import-Normal\"><strong>Monogamy<\/strong>: A mating system in which one male mates with one female.<\/p>\n<p class=\"import-Normal\"><strong>Multi-male, multi-female<\/strong>: A group that consists of multiple adult males, multiple adult females, and their dependent offspring.<\/p>\n<p class=\"import-Normal\"><strong>Multi-male, single-female<\/strong>: A group that consists of two or more adult males, one breeding female, their dependent offspring, and non-breeding females.<\/p>\n<p class=\"import-Normal\"><strong>Mutualistic\/mutualism<\/strong>: When different species work together, with each benefiting from the interaction.<\/p>\n<p class=\"import-Normal\"><strong>Niche<\/strong>: The role of a species in its environment; how it meets its needs for food, shelter, etc.<\/p>\n<p class=\"import-Normal\"><strong>Nocturnal<\/strong>: Active at night.<\/p>\n<p class=\"import-Normal\"><strong>Olfactory communication<\/strong>: Conveying information through scent.<\/p>\n<p class=\"import-Normal\"><strong>Omnivores<\/strong>: Organisms whose diet consists of plant and animal matter.<\/p>\n<p class=\"import-Normal\"><strong>Pair bond<\/strong>: A strong, long-term relationship between two individuals.<\/p>\n<p class=\"import-Normal\"><strong>Parasite<\/strong>: An organism that lives in or on another organism.<\/p>\n<p class=\"import-Normal\"><strong>Parental investment<\/strong>: Any time or energy a parent devotes to the current offspring that enhances its survival (and eventual reproductive success) at the expense of the parent\u2019s ability to invest in the next offspring.<\/p>\n<p class=\"import-Normal\"><strong>Philopatric<\/strong>: Remaining in the group of one\u2019s birth.<\/p>\n<p class=\"import-Normal\"><strong>Piloerection<\/strong>: Raising one\u2019s hair or fur in an effort to look bigger.<\/p>\n<p class=\"import-Normal\"><strong>Polyandry<\/strong>: A mating system in which multiple males mate with a single breeding female.<\/p>\n<p class=\"import-Normal\"><strong>Polygamy<\/strong>: A mating system in which multiple males mate with multiple females.<\/p>\n<p class=\"import-Normal\"><strong>Polygyny<\/strong>: A mating system in which one male mates with multiple females.<\/p>\n<p class=\"import-Normal\"><strong>Polyspecific association<\/strong>: An association between two or more different species that involves behavioural changes in at least one of them to maintain the association.<\/p>\n<p class=\"import-Normal\"><strong>Primate community<\/strong>: All primate species that occur in an area.<\/p>\n<p class=\"import-Normal\"><strong>Primatologist<\/strong>: A scientist who studies primate behaviour and\/or ecology.<\/p>\n<p class=\"import-Normal\"><strong>Primatology<\/strong>: The scientific field that studies primate behaviour and\/or ecology.<\/p>\n<p class=\"import-Normal\"><strong>Ranging behaviour<\/strong>: Refers to the way in which animals move about their environment.<\/p>\n<p class=\"import-Normal\"><strong>Receptive<\/strong>: A term used to describe females who are ready for sexual reproduction (i.e., not pregnant or nursing).<\/p>\n<p class=\"import-Normal\"><strong>Reproductive success<\/strong>: An individual\u2019s genetic contribution to future generations, often measured through the number of offspring produced.<\/p>\n<p class=\"import-Normal\"><strong>Reproductive suppression<\/strong>: The prevention or inhibition of reproduction of healthy adults.<\/p>\n<p class=\"import-Normal\"><strong>Resident male<\/strong>: Term that describes the male who lives with a group of females.<\/p>\n<p class=\"import-Normal\"><strong>Seed dispersal<\/strong>: The process by which seeds move away from the plant that produced them in preparation for germination and becoming a new plant.<\/p>\n<p class=\"import-Normal\"><strong>Semantic communication<\/strong>: The systematic use of signals to refer to objects in the environment.<\/p>\n<p class=\"import-Normal\"><strong>Sexual dimorphism<\/strong>: When males and females of a species have different morphological traits.<\/p>\n<p class=\"import-Normal\"><strong>Sexual selection<\/strong>: The selection for traits that increase mating success. This occurs via intersexual selection and intrasexual selection.<\/p>\n<p class=\"import-Normal\"><strong>Sexual swelling<\/strong>: Area of the hindquarters that change in size, shape, and often colour over the course of a female\u2019s reproductive cycle, reaching maximum size at ovulation. Occurs in many primate species that live in Africa and Asia.<\/p>\n<p class=\"import-Normal\"><strong>Sexually monomorphic<\/strong>: When males and females of a species have similar morphological traits.<\/p>\n<p class=\"import-Normal\"><strong>Single-male, multi-female<\/strong>: A group that consists of one adult male, multiple adult female, and their dependent offspring.<\/p>\n<p class=\"import-Normal\"><strong>Single-male, single-female<\/strong>: A group that consists of one adult male, one adult female, and their dependent offspring.<\/p>\n<p class=\"import-Normal\"><strong>Social learning<\/strong>: The idea that new behaviours can be acquired by observing and imitating others.<\/p>\n<p class=\"import-Normal\"><strong>Social system<\/strong>: A way of describing the typical number of males and females of all age classes that live together.<\/p>\n<p class=\"import-Normal\"><strong>Social transmission<\/strong>: Transfer of something from one individual to another; this can include parasites, information, or cultural traditions.<\/p>\n<p class=\"import-Normal\"><strong>Sociality<\/strong>: The tendency to form social groups.<\/p>\n<p class=\"import-Normal\"><strong>Solitary<\/strong>: Living alone.<\/p>\n<p class=\"import-Normal\"><strong>Species recognition<\/strong>: The ability to differentiate conspecifics from members of other species.<\/p>\n<p class=\"import-Normal\"><strong>Subordinate<\/strong>: Being of low rank.<\/p>\n<p class=\"import-Normal\"><strong>Tactile communication<\/strong>: Conveying information through touch.<\/p>\n<p class=\"import-Normal\"><strong>Territory:<\/strong> A home range whose boundary is defended from intrusion by conspecifics.<\/p>\n<p class=\"import-Normal\"><strong>Vertebrates<\/strong>: The group of animals characterized by an internal spinal column or backbone. This includes fish, amphibians, reptiles, birds, and mammals.<\/p>\n<p class=\"import-Normal\"><strong>Vigilance<\/strong>: Watchful behaviour used to detect potential danger, usually in the form of predators or potential competitors.<\/p>\n<p class=\"import-Normal\"><strong>Visual communication<\/strong>: Conveying information through signals that can be seen.<\/p>\n<p class=\"import-Normal\"><strong>Vocal communication<\/strong>: Conveying information through signals that can be heard.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration<\/h2>\n<p class=\"import-Normal\">Goodall, Jane. 1971. <em>In the Shadow of Man<\/em>. Boston: Houghton Mifflin.<\/p>\n<p class=\"import-Normal\">Rowe, Noel, and Marc Myers, eds. 2016. <em>All the World\u2019s Primates. <\/em>Charleston, RI: Pogonias Press.<\/p>\n<p class=\"import-Normal\">Strier, Karen B. 2017. <em>Primate Behavioural Ecology.<\/em> 5th ed. New York: Routledge.<\/p>\n<p class=\"import-Normal\"><a href=\"https:\/\/pin.primate.wisc.edu\/\">Primate Info Net<\/a> is an information service of the National Primate Research Center at the University of Wisconsin, Madison. It includes Primate Factsheets, primate news and publications, a list of primate-related jobs, and an international directory of primatology, among other information.<\/p>\n<p class=\"import-Normal\"><a href=\"https:\/\/www.primate-sg.org\/\">Primate Specialist Group<\/a> is a collection of scientists and conservationists who work in dozens of African, Asian, and Latin American nations to promote research on primate conservation.<\/p>\n<p class=\"import-Normal\">Short videos of some primate behaviours discussed in this chapter:<\/p>\n<ul>\n<li class=\"import-Normal\">Watch vervet monkeys respond to different types of predators: BBC One. n.d. \u201cVervet Monkey\u2019s Escape Plans - Talk to the Animals: Episode 2 Preview.\u201d Accessed December 16, 2022. <a class=\"rId10\" href=\"https:\/\/www.youtube.com\/watch?v=q8ZG8Dpc8mM\">https:\/\/www.youtube.com\/watch?v=q8ZG8Dpc8mM. <\/a><\/li>\n<li class=\"import-Normal\">Watch male gelada baboons use the lip flip in competition with other males: Smithsonian Channel, June 9, 2017. \u201cWhy These Vegetarian Monkeys Have Sharp Predator Teeth.\u201d Accessed July 25, 2019. <a class=\"rId11\" href=\"https:\/\/www.youtube.com\/watch?time_continue=145&amp;v=aC6iYj_EBjY\">https:\/\/www.youtube.com\/watch?time_continue=145&amp;v=aC6iYj_EBjY<\/a>.<\/li>\n<li class=\"import-Normal\">Watch (and listen to!) howler monkeys \u201croar\u201d: Science News. N.d. \u201cHear a Male Howler Monkey Roar.\u201d Accessed November 21, 2022. <a class=\"rId12\" href=\"https:\/\/www.youtube.com\/watch?v=PYar0dkZ6v8\">https:\/\/www.youtube.com\/watch?v=PYar0dkZ6v8<\/a>.<\/li>\n<li class=\"import-Normal\">Watch Japanese macaques using natural hot springs: National Geographic. N.d. \u201cMeditative Snow Monkeys Hang Out in Hot Springs.\u201d Accessed July 25, 2019. <a class=\"rId13\" href=\"https:\/\/www.youtube.com\/watch?v=Aat9O85ynsI\">https:\/\/www.youtube.com\/watch?v=Aat9O85ynsI<\/a>.<\/li>\n<li class=\"import-Normal\">Watch chimpanzees make and use tools: National Geographic. n.d. \u201cChimps and Tools.\u201d Accessed July 25, 2019. <a class=\"rId14\" href=\"https:\/\/www.youtube.com\/watch?v=o2TBicMRLtA\">https:\/\/www.youtube.com\/watch?v=o2TBicMRLtA<\/a>.<\/li>\n<\/ul>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\">Aich, H., R. Moos-Heilen, and E. Zimmermann. 1990. \u201cVocalizations of Adult Gelada Baboons (<em>Theropithecus gelada<\/em>): Acoustic Structure and Behavioural Context.\u201d <em>Folia Primatologica<\/em> 55 (3\u20134): 109\u2013132.<\/p>\n<p class=\"import-Normal\">Bell, Sarah A. 2017. \u201cGaldikas, Birute.\u201d In <em>The International Encyclopedia of Primatology, Volume A\u2013G<\/em>, edited by Agust\u00edn Fuentes, 445\u2013446. Malden, MA: John Wiley &amp; Sons.<\/p>\n<p class=\"import-Normal\">Boinski, S. 1992. \u201cOlfactory Communication among Costa Rican Squirrel Monkeys: A Field Study.\u201d <em>Folia Primatologica<\/em> 59 (3): 127\u2013136.<\/p>\n<p class=\"import-Normal\">Cheney, D. L., and R. M. Seyfarth. 1987. \u201cThe Influence of Intergroup Competition on the Survival and Reproduction of Female Vervet Monkeys.\u201d <em>Behavioral Ecology and Sociobiology<\/em> 21 (6): 375\u2013386.<\/p>\n<p class=\"import-Normal\">de Oliveira Terceiro, Francisco Edvaldo, and Judith M. Burkart. 2019. \u201cCooperative Breeding.\u201d In <em>Encyclopedia of Animal Cognition and Behavior<\/em>, edited by Jennifer Vonk and Todd Shackelford, 1\u20136. Edinburg, Scotland: Springer Cham.<\/p>\n<p class=\"import-Normal\">Digby, Leslie J., Stephen F. Ferrari, and Wendy Saltzman. 2011. \u201cCallitrichines: The Role of Competition in Cooperatively Breeding Species.\u201d In <em>Primates in Perspective<\/em>, edited by Christina J. Campbell, August\u00cdn Fuentes, Katherine C. MacKinnon, Simon K. Bearder, and Rebecca M. Stumpf, 91\u201310. 2nd edition. New York: Oxford University Press.<\/p>\n<p class=\"import-Normal\">Fischer, Julia, Kurt Hammerschmidt, Dorothy L. Cheney, and Robert M. Seyfarth. 2008. \u201cAcoustic Features of Female Chacma Baboon Barks.\u201d <em>Ethology<\/em> 107 (1): 33\u201354.<\/p>\n<p class=\"import-Normal\">Jolly, Alison. 1966. <em>Lemur Behavior: A Madagascar Field Study<\/em>. Chicago: University of Chicago Press.<\/p>\n<p class=\"import-Normal\">Krief, Sabrina, Claude Marcel Hladik, and Claudie Haxaire. 2005. \u201cEthnomedicinal and Bioactive Properties of Plants Ingested by Wild Chimpanzees in Uganda.\u201d <em>Journal of Ethnopharmacology<\/em> 110 (1\u20133): 1\u201315.<\/p>\n<p class=\"import-Normal\">Maekawa, Mkio, Annette Lanjouw, Eug\u00e8ne Rutagarama, and Doublas Sharp. 2013. \u201cMountain Gorilla Tourism Generating Wealth and Peace in Post-Conflict Rwanda.\u201d <em>Natural Resources Forum<\/em> 37 (2): 127\u2013137.<\/p>\n<p class=\"import-Normal\">Matsuzawa, Tetsuro. 2015. \u201cSweet-Potato Washing Revisited: 50th Anniversary of the <em>Primates<\/em> Article.\u201d <em>Primates<\/em> 56: 285\u2013287.<\/p>\n<p class=\"import-Normal\">Matsuzawa, Tetsuro. 2018. \u201cHot-Spring Bathing of Wild Monkeys in Shiga-Heights: Origin and Propagation of a Cultural Behavior.\u201d <em>Primates<\/em> 59: 209\u2013213.<\/p>\n<p class=\"import-Normal\">McGrew, W. C. 1998. \u201cCulture in Nonhuman Primates?\u201d <em>Annual Review of Anthropology<\/em> 27: 301\u2013328.<\/p>\n<p class=\"import-Normal\">Mertl-Millhollen, Anne S. 1988. \u201cOlfactory Demarcation of Territorial but Not Home Range Boundaries by <em>Lemur catta<\/em>.\u201d <em>Folia Primatologica<\/em> 50 (3\u20134): 175\u2013187.<\/p>\n<p class=\"import-Normal\">Pinacho-Guendulain, B., and G. Ramos-Fern\u00e1ndez. 2017. \u201cInfluence of Fruit Availability on the Fission-Fusion Dynamics of Spider Monkeys (<em>Ateles geoffroyi<\/em>).\u201d <em>International Journal of Primatology<\/em> 38: 466\u2013484.<\/p>\n<p class=\"import-Normal\">Poirotte, Cl\u00e9mence, Fran\u00e7ois Massol, Ana\u00efs Herbert, Eric Willaume, Pacelle M. Bomo, Peter M. Kappeler, and Marie J. E. Charpentier. 2017. \u201cMandrills Use Olfaction to Socially Avoid Parasitized Conspicifics.\u201d <em>Science Advances<\/em> 3 (4): e160172.<\/p>\n<p class=\"import-Normal\">Rodrigues, Michelle. 2019. \u201cIt\u2019s Time to Stop Lionizing Dian Fossey as a Conservation Hero.\u201d <em>Lady Science<\/em> website, September 20. Accessed December 14, 2022. <a class=\"rId15\" href=\"https:\/\/www.ladyscience.com\/ideas\/time-to-stop-lionizing-dian-fossey-conservation\">https:\/\/www.ladyscience.com\/ideas\/time-to-stop-lionizing-dian-fossey-conservation<\/a>.<\/p>\n<p class=\"import-Normal\">Samuni, Liran, Anna Preis, Tobias Deschner, Catherine Crockford, and Roman M. Wittig. 2018. \u201cReward of Labor Coordination and Hunting Success in Wild Chimpanzees.\u201d <em>Communications Biology<\/em> 1: 138.<\/p>\n<p class=\"import-Normal\">Santana, Sharlene E., Jessica Lynch Alfaro, and Michael E. Alfaro. 2012. \u201cAdaptive Evolution of Facial Colour Patterns in Neotropical Primates.\u201d <em>Proceedings of the Royal Society B: Biological Sciences<\/em> 279 (1736): 2204\u20132211.<\/p>\n<p class=\"import-Normal\">Sanz, Crickette M., David Strait, Crepin Eyana Ayina, Jean Marie Massamba, Thierry Fabrice Ebombi, Severin Ndassoba Kialiema, Delon Ngoteni, et al. 2022. \u201cInterspecific Interactions Between Sympatric Apes.\u201d i<em>Science<\/em> 25 (10): 105059.<\/p>\n<p class=\"import-Normal\">Sch\u00f6n Ybarra, M. A. 1986. \u201cLoud Calls of Adult Male Red Howling Monkeys (<em>Alouatta seniculus<\/em>).\u201d <em>Folia Primatologica<\/em> 47 (4): 204\u2013216.<\/p>\n<p class=\"import-Normal\">Setchell, Joanna M., Tessa Smith, E. Jean Wickings, and Leslie A. Knapp. 2008. \u201cSocial Correlates of Testosterone and Ornamentation in Male Mandrills.\u201d <em>Hormones and Behavior<\/em> 54 (3): 365\u2013372.<\/p>\n<p class=\"import-Normal\">Setchell, Joanna M., E. Jean Wickings, and Leslie A. Knapp. 2006. \u201cSignal Content of Red Facial Coloration in Female Mandrills (<em>Mandrillus sphinx<\/em>).\u201d <em>Proceedings of the Royal Society B: Biological Sciences<\/em><a class=\"rId16\" href=\"https:\/\/paperpile.com\/b\/Gb7Zko\/Lxpl\"> 273 (1599): 2395\u20132400.<\/a><\/p>\n<p class=\"import-Normal\">Seyfarth, R. M., D. L. Cheney, and P. Marler. 1980a. \u201cMonkey Responses to Three Different Alarm Calls: Evidence of Predator Classification and Semantic Communication.\u201d <em>Science<\/em> 210 (4471): 801\u2013803.<\/p>\n<p class=\"import-Normal\">Seyfarth, Robert M., Dorothy L. Cheney, and Peter Marler. 1980b. \u201cVervet Monkey Alarm Calls: Semantic Communication in a Free-Ranging Primate.\u201d <em>Animal Behaviour<\/em> 28 (4): 1070\u20131094.<\/p>\n<p class=\"import-Normal\">Sharma, Goutam, Chan Ram, and Lal Singh Rajpurohit. 2010. \u201cA Case Study of Infantcide After Resident Male Replacement in <em>Semnopithecus entellus<\/em> around Jodhpur (India).\u201d <em>Proceeding of the Zoological Society<\/em> 63 (2): 93\u201398.<\/p>\n<p class=\"import-Normal\">Shirasu, Mika, Satomi Ito, Akihiro Itoigawa, Takashi Hayakawa, Kodzue Kinoshita, Isao Munechika, Hiroo Imai, and Kazushige Touhara. 2020. \u201cKey Male Glandular Odorants Attracting Female Ring-Tailed Lemurs.\u201d <em>Current Biology<\/em> 30 (11): 2131\u20132138.<\/p>\n<p class=\"import-Normal\">Stanford, Craig B. 2017. \u201cGoodall, Jane.\u201d In <em>The International Encyclopedia of Primatology, Volume A\u2013G<\/em>, edited by Agust\u00edn Fuentes, 471\u2013472. Malden, MA: John Wiley &amp; Sons.<\/p>\n<p class=\"import-Normal\">Stewart, Kelly. 2017. \u201cFossey, Dian.\u201d In <em>The International Encyclopedia of Primatology, Volume A\u2013G<\/em>, edited by Agust\u00edn Fuentes, 432\u2013433. Malden, MA: John Wiley &amp; Sons.<\/p>\n<p class=\"import-Normal\">Takeshita, Rafaela S.C., Fred B. Bercovitch, Kodzue Kinoshita, and Michael A. Huffman. 2018. \u201cBeneficial Effect of Hot Spring Bathing on Stress Levels in Japanese Macaques.\u201d <em>Primates<\/em> 59 (3): 215\u2013225.<\/p>\n<p class=\"import-Normal\">Trivers, Robert L. 1972. \u201cParental Investment and Sexual Selection.\u201d In <em>Sexual Selection and the Descent of Man, 1871\u20131971<\/em>, edited by Bernard Campbell, 136\u2013179. Chicago: Aldine.<\/p>\n<p class=\"import-Normal\">Whiten, Andrew. 2011. \u201cThe Scope of Culture in Chimpanzees, Humans and Ancestral Apes.\u201d <em>Philosophical Transactions of the Royal Society of London B: Biological Sciences<\/em> 366 (1567): 997\u20131007.<\/p>\n<p class=\"import-Normal\">Wiens, Frank, and Annette Zitzmann. 2003. \u201cSocial Structure of the Solitary Slow Loris <em>Nycticebus coucang<\/em> (Lorisidae).\u201d <em>Journal of Zoology<\/em> 261 (1): 35\u201346.<\/p>\n<p class=\"import-Normal\">Zuberb\u00fchler, Klaus, David Jenny, and Redouan Bshary. 1999. \u201cThe Predator Deterrence Function of Primate Alarm Calls.\u201d <em>Ethology<\/em> 105 (6): 477\u2013490.<\/p>\n<p class=\"import-Normal\">Zuberb\u00fchler, Klaus, Ronald No\u00eb, and Robert M. Seyfarth. 1997. \u201cDiana Monkey Long-Distance Calls: Messages for Conspecifics and Predators.\u201d <em>Animal Behaviour<\/em> 53 (3): 589\u2013604.<\/p>\n<h2 class=\"import-Normal\">Acknowledgements<\/h2>\n<p class=\"import-Normal\">The author is grateful to the editors for the opportunity to contribute to this open-source textbook. She thanks Dr. Stephanie Etting for her encouragement and support during the revision of this chapter. Her suggestions, along with comments made by two anonymous reviewers on an earlier draft of this chapter, improved the final version considerably. Finally, she thanks all the primatologists who came before her, especially her advisor, Lynne A. Isbell, for their tireless efforts to understand the behavior and ecology of the living primates. Without their work, this chapter would not have been possible.<\/p>\n<\/div>\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_1683_1711\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1711\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<div class=\"textbox\">\n<h6>Student contributors for this chapter from Salinda Hess' ANTH 203: <em>Emma Costa, Shima Gahima, Will Lefebvre, Audrey Ch\u00e9kina\u00ebl. <\/em><\/h6>\n<p>Learn more about <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/about-the-authors\/\">authors and editors<\/a>,<a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/student-contributors\/\"> Hess' student contributions<\/a>, and <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/versioning-history\/\">versioning history<\/a>.<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\">Learning Objectives<\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li class=\"import-Normal\">Describe the variables that affect primate diets.<\/li>\n<li class=\"import-Normal\">Explain how primates interact with other organisms in their environment.<\/li>\n<li class=\"import-Normal\">Discuss why primates live in groups, types of primate groups, and components of their social systems.<\/li>\n<li class=\"import-Normal\">Describe the reproductive strategies of males and females.<\/li>\n<li class=\"import-Normal\">Explain the ways in which primates communicate.<\/li>\n<li class=\"import-Normal\">Discuss the evidence for primate cultural traditions.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p class=\"import-Normal\">Nonhuman primates (hereafter, 'primates') are a fascinating group of animals, whose similarity to humans can be striking. Because of this similarity, studying primates helps anthropologists to gain insight into how our human ancestors may have behaved. It also allows us to better understand our own behaviour through <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_928\">comparison<\/a><\/strong> (examining similarities and differences) with other primates as well as by comparing different species of primates to one another. In this way, studying primates helps anthropologists comprehend humanity from a biological perspective, which contributes to anthropology\u2019s commitment to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_930\">holism,<\/a><\/strong> the idea that the parts of a system interconnect and interact to make up the whole.<\/p>\n<figure style=\"width: 242px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2023\/05\/image3.png\" alt=\"A person using binoculars to look at monkeys.\" width=\"242\" height=\"354\" \/><figcaption class=\"wp-caption-text\">Figure 7.1: The author observing patas monkeys from a distance in Laikipia, Kenya. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-6\/\">Karin Enstam Jaffe observing patas monkeys in Laikipia, Kenya (Figure 6.5)<\/a> by Rebecca Chancellor is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_932\">Ethology<\/a> <\/strong>is the study of animal behaviour, while <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_934\">primatology <\/a><\/strong>is the study of primate behaviour. People who study primates are called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_936\">primatologists<\/a><\/strong>. Research on primates can be conducted in the field (i.e., on wild primates) or in captivity (i.e., zoos) and may or may not involve experiments, such as playing recorded alarm calls to see how individuals react. Unlike some other Science, Technology, Engineering, and Math (STEM) fields, primatology has a long history of research conducted by women (see \u201cSpecial Topic: Women in Primatology\u201d). Primatologists come from many different disciplines, have diverse backgrounds, and study primates for different reasons. Biologists study primates as examples of evolutionary theories like natural selection, and to understand behaviours as <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_938\">adaptations<\/a><\/strong>, or traits with a function that increases <strong>fitness<\/strong>, i.e. an individual\u2019s survival and\/or reproduction. Primate intelligence is of interest to psychologists who want to learn more about deception or cooperation and to linguists interested in the principles of communication and language. Ecologists consider how primates interact with the habitats they occupy, and conservationists examine how primates are affected by deforestation, poaching, or illegal animal trade (see Appendix B: Primate Conservation for more information on these topics). Biological anthropologists, like myself (Figure 7.1), who study primates are interested in learning about their social complexity, and ecological and behavioural variation, to better understand the biological basis of human behaviour. And, similar to biologists, we also explore how primate behaviour is adaptive and contributes to individual fitness. Like other sciences, primatology is only as strong as its researchers, methods, and theories, and the field has benefitted recently from efforts to increase diversity and reckon with its colonialist past, as discussed below in \u201cSpecial Topic: Women in Primatology.\u201d<\/p>\n<p class=\"import-Normal\">Humans share many traits in common with primates. As you learned in Chapter 5, some of these traits are similar due to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_944\">homology<\/a><\/strong>, traits both species inherited from a common primate ancestor. For example, like most other primates, humans are social animals who live in groups. Group living did not evolve independently in humans and other primates. Rather, group living is a trait that evolved in a primate ancestor, and because it benefited survival, it was retained in the species\u2019 <strong>descendants<\/strong> (or the species that come after the ancestor species). In contrast, humans and other primates can have similar traits that evolved independently, which is called <strong>analogy<\/strong>. For example, both humans and Japanese macaques (<em>Macaca fuscata<\/em>) use natural hot springs (Figures 7.2a &amp; b). Research on these monkeys indicates that sitting in hot springs reduces stress and helps keep them warm, much as it does for humans (Takeshita Et al. 2018). But this behaviour is not the result of humans and Japanese macaques having a shared ancestor who used hot springs. Rather, the behaviour arose independently in two species that both occupy northerly environments and adapted to cold climates using a similar behaviour. Studying the homologous traits we share with other primates, like living in groups, helps us develop hypotheses about human behaviours as adaptations, which in turn helps us develop models for the behaviour of our human ancestors. Studying analogous traits, like hot springs use, allows us to better understand the effects of ecological variables on morphology and behaviour of both primates and humans, living and extinct.<\/p>\n<figure id=\"attachment_191-5\" aria-describedby=\"caption-attachment-191-5\" style=\"width: 2161px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-189 size-full\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.2.jpg\" alt=\"Left, a man in a hot spring. Right, monkeys in a hot spring.\" width=\"2161\" height=\"803\" \/><figcaption id=\"caption-attachment-191-5\" class=\"wp-caption-text\">Figure 7.2a\/b: Both humans (left) and Japanese macaques (right) use natural hot springs to reduce stress and relax. This similar trait arose independently in the two species, making it a good example of analogy. Credit: 7.2a. <a href=\"https:\/\/pixabay.com\/photos\/hot-spring-landscape-man-mountain-1846721\/\">Hot Spring Landscape<\/a> by <a href=\"https:\/\/pixabay.com\/users\/pexels-2286921\/?utm_source=link-attribution&amp;utm_medium=referral&amp;utm_campaign=image&amp;utm_content=1846721\">Pexels<\/a> has been modified (cropped) and 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>. 7.2b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Jigokudani_hotspring_in_Nagano_Japan_001.jpg\">Jigokudani hotspring in Nagano Japan 001<\/a> by Yosemite 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<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Women in Primatology<\/h2>\n<p class=\"import-Normal\">While many STEM fields have traditionally been, and continue to be, dominated by men, primatology has a long history of significant research conducted by women. This is due, in part, to the fact that three of the most well-known primatologists are women. In the early 1960s, British paleoanthropologist Louis Leakey (discussed in Chapters 9 and 10) was looking for students to study the great apes in hopes of shedding light on the behaviours of our early ancestors. He chose Jane Goodall (Figure 7.3a) to study chimpanzees (<em>Pan troglodytes<\/em>), Birute Galdikas (Figure 7.3b) to study Bornean orangutans (<em>Pongo pygmaeus<\/em>), and Dian Fossey (Figure 7.3c) to study mountain gorillas (<em>Gorilla<\/em><em> beringei beringei<\/em>). The work of these three women, sometimes referred to as Leakey\u2019s \u201cTrimates,\u201d has transformed our understanding of ape (and primate) behaviour.<\/p>\n<figure id=\"attachment_190-3\" aria-describedby=\"caption-attachment-190-3\" style=\"width: 626px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-190\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.3-1.jpg\" alt=\"Jane Goodall, Birute Galdikas, and Dian Fossey.\" width=\"626\" height=\"209\" \/><figcaption id=\"caption-attachment-190-3\" class=\"wp-caption-text\">Figure 7.3a-c: Louis Leakey\u2019s \u201cTrimates\u201d (left to right): a. Jane Goodall\u2019s research on the Gombe chimpanzees spans over half a century; b. Birute Galdikas\u2019s research and rescue work on behalf of orangutans spans 40 years; c. Dian Fossey studied mountain gorillas in Rwanda for almost 20 years, until her murder in 1985. Credit: a. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Jane_Goodall_HK.jpg\">Jane Goodall HK<\/a> by Jeekc has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\"> CC BY-SA 3.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Dr_Birute_Galdikas.jpg\">Dr Birute Galdikas<\/a> by<a href=\"https:\/\/www.flickr.com\/photos\/sfupamr\/\"> Simon Fraser University - University Communications<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\"> CC BY 2.0 License<\/a>. c. <a href=\"https:\/\/www.flickr.com\/photos\/mary-lynn\/2925879356\">US-223658 Dian Fossey<\/a> by<a href=\"https:\/\/www.flickr.com\/photos\/mary-lynn\/2925879356\/\"> Mary-Lynn<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\"> CC BY 2.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Arriving at the Gombe Stream Reserve in Tanzania in 1960, Jane Goodall was one of the first scientists to conduct a long-term study of wild nonhuman primates. Before then, most studies lasted less than a year and were often zoo-based. By 1961, she had made two astounding observations that forced us to reconsider what differentiates humans from the rest of the primate order. She observed chimpanzees eating a colobus monkey, the first reported evidence of meat eating in our closest relatives; later observing them hunting and sharing meat. And she discovered that chimpanzees make and use tools by stripping leaves off twigs to \u201cfish\u201d for termites. Her work, spanning several decades, has produced long-term data on chimpanzee mating strategies, mother-infant bonds, and aggression. In the mid-1980s, Goodall transitioned from field researcher to conservationist and activist, advocating for the humane use of nonhuman animals (Stanford 2017).<\/p>\n<p class=\"import-Normal\">Birute Galdikas began her study of orangutans in Kalimantan, Borneo, in 1971. Hers was the first long-term study conducted on the Bornean orangutan. Galdikas and her colleagues have collected over 150,000 hours of observational data, focusing on the life histories of individual orangutans. While conducting behavioural research, Galdikas discovered that the pet trade and habitat loss were adversely affecting the orangutan population. Eventually, Galdikas\u2019s conservation efforts began to extend beyond advocacy and into rehabilitation and forest preservation (Bell 2017). If you would like to learn more about primate conservation efforts, please see Appendix B: Primate Conservation.<\/p>\n<p class=\"import-Normal\">In 1967, Dian Fossey began her long-term study of mountain gorillas and founded the Karisoke Research Centre in Rwanda. Her and her colleagues\u2019 research, over several decades, revealed much about gorilla social behaviour, ecology, and life history. Her efforts also led to the development of mountain gorilla conservation programs. However, she was a controversial figure, as discussed below. Fossey was murdered in December 1985; the case remains unsolved (Stewart 2017).<\/p>\n<h3 class=\"import-Normal\"><strong>Decolonizing Primatology<\/strong><\/h3>\n<p class=\"import-Normal\">Recently, the movement to <strong>decolonize<\/strong> primatology, by understanding and highlighting the theories and research of non-Western individuals and perspectives, has gathered steam. This movement draws attention to the maltreatment of local people by Western primatologists. For example, Michelle Rodrigues (2019) argues that it's time we stop focusing on the scientific and conservation contributions of Dian Fossey and acknowledge that her \"active conservation\" techniques included kidnapping and torturing local Rwandans who were known as, or suspected to be, gorilla poachers. Rodrigues (2019) argues:<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 36pt;text-indent: 0pt\">The image of Fossey, a white American woman, whipping and torturing black African poachers is evocative of the behaviour of white slaveholders in the American South. It is appalling enough to think of that behaviour occurring in the 1850s; there is no way we can explain Fossey\u2019s behaviour in the 1970s as the product of \u201ca different time.\u201d Yet, almost three decades later, the romantic notion of a noble martyr who died for her devotion to gorillas prevails, and these terrifying actions are often described as simply unorthodox methods. Perhaps these truths are softened due to fears that the reality of this legacy would harm gorilla conservation efforts. But memorializing her as a martyr and patron saint of gorilla conservation demands that we forget the cruel acts she advocated for and performed.<\/p>\n<p class=\"import-Normal\">Further, Louis Leakey\u2019s installment of Goodall, Galdikas, and Fossey to study chimpanzees, orangutans, and mountain gorillas, respectively, is itself viewed as recapitulating the colonial legacy in Africa and Asia. Given that Leakey was the offspring of British missionaries, Rodrigues (2019) argues, it is no accident that he was willing to mentor British and American women, while overlooking women from Africa and Asia as potential researchers. This leads us to another level of the decolonizing movement, which aims to highlight the research of non-Western primatologists, particularly those living in what primatologists refer to as \u201chabitat countries\u201d that are home to living primates. As you will see in this chapter, scientists from diverse backgrounds are active contributors to exciting research on primates around the world.<\/p>\n<\/div>\n<h2 class=\"import-Normal\">Ecology<\/h2>\n<p class=\"import-Normal\">The more than 600 species and subspecies of living primates are highly diverse in their dietary preferences and the habitats they occupy. In this section we\u2019ll briefly discuss aspects of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_950\">ecology<\/a><\/strong>, or the relationship between organisms and their physical surroundings, that impact a primate\u2019s life, the foods they eat, and the other species with whom they interact.<\/p>\n<h3 class=\"import-Normal\"><strong>Primate Diets<\/strong><\/h3>\n<p class=\"import-Normal\">Diet may be the most important variable influencing variation in primate morphology, behaviour, and ecology. Most primates are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_952\">omnivores<\/a><\/strong> who ingest a variety of foods in order to obtain appropriate levels of protein, carbohydrates, fats, and fluids, but one type of food often makes up the majority of each species\u2019 diet. You learned about the dental and digestive adaptations of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_954\">frugivores<\/a><\/strong> (who feed primarily on fruit), <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_956\">folivores<\/a><\/strong> (whose diet consists mostly of leaves), and <strong>insectivores <\/strong>(who eat mainly insects) in Chapter 6, so we will not discuss them again here.<\/p>\n<h4 class=\"import-Normal\"><em>Body Size and Diet<\/em><\/h4>\n<figure id=\"attachment_191-6\" aria-describedby=\"caption-attachment-191-6\" style=\"width: 559px\" class=\"wp-caption alignright\"><img class=\"wp-image-191\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.4-1.jpg\" alt=\"A tarsier eats a grasshopper. A gorilla eats leaves.\" width=\"559\" height=\"237\" \/><figcaption id=\"caption-attachment-191-6\" class=\"wp-caption-text\">Figure 7.4a-b: Primates eat different types of food. Small primates, like the spectral tarsier (left), eat mostly insects while large primates, like the mountain gorilla (right), eat mostly leaves. Credit: a. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Spectral_Tarsier_Tarsius_tarsier_(7911549768).jpg\">Spectral Tarsier Tarsius tarsier (7911549768)<\/a> by<a href=\"https:\/\/www.flickr.com\/people\/65695019@N07\"> Bernard DUPONT<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/legalcode\"> CC BY-SA 2.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Mountain_gorilla_(Gorilla_beringei_beringei)_eating.jpg\">Mountain gorilla (Gorilla beringei beringei) eating<\/a> by<a href=\"https:\/\/www.sharpphotography.co.uk\/\"> Charles J Sharp<\/a> (creator QS:P170,Q54800218) has been modified (cropped) 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>Insects are a high-quality food, full of easily digestible protein and high in calories that meet most of a primate\u2019s dietary needs. Although all primates will eat insects if they come upon them, those species that rely most heavily on insects tend to be the smallest. Why? Because larger primates simply cannot capture and consume enough insects every day to survive. Because of their small size (less than 150 g), spectral tarsiers (<em>Tarsius spectrum<\/em>) have a fast <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_960\">metabolism<\/a><\/strong>, which means they turn food to energy quickly, but they do not need to consume large amounts of food each day. It does not matter to a spectral tarsier that a grasshopper only weighs 300 mg, because the tarsier (<em>Tarsius<\/em>) itself is so small that one grasshopper is a good-size meal (Figure 7.4a). That same grasshopper is not even a snack for an adult male mountain gorilla (<em>Gorilla beringei beringei<\/em>), who may weigh up to 200 kg. Fortunately for gorillas (<em>Gorilla)<\/em>, their large body size means they have a slow metabolism, converting food into energy much more slowly, so they can eat lower quality food that takes longer to digest, provided there is a lot of it. For gorillas, leaves, which are hard to digest but plentiful, fit the bill (Figure 7.4b). Most medium-sized primates are highly frugivorous, and supplement their fruit based diet in ways that correspond with their size: Smaller frugivores tend to supplement with insects, while larger frugivores tend to supplement with leaves.<\/p>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<h4 class=\"import-Normal\"><em>Food Abundance and Distribution<\/em><\/h4>\n<p class=\"import-Normal\">Nutrients are not the only dietary considerations primates must make. They must also ensure that they consume more calories than they use. The abundance and distribution of food affect energy expenditure and calorie intake because they determine how far animals must travel in search of food and how much they must compete to obtain it. <strong>Abundance <\/strong>refers to how much food is available in a given area while <strong>distribution<\/strong> refers to how food is spread out. In terms of abundance, food is either plentiful or scarce (Figure 7.5a\u2013b). Food is distributed in one of three ways: uniformly (Figure 7.6a), in clumps (Figure 7.6b), or randomly (Figure 7.6c). In general, higher-quality foods, like fruit and insects, are less abundant and have patchier distributions than lower-quality foods, like leaves. Primates who eat fruit or insects usually have to travel farther to find food and burn more calories in the process. Abundance and distribution of food is another reason why larger primates tend to rely more heavily on leaves than either fruit or insects.<\/p>\n<figure style=\"width: 619px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image2-2.png\" alt=\"Two squares with different amounts of dots. \" width=\"619\" height=\"286\" \/><figcaption class=\"wp-caption-text\">Figure 7.5a-b: Two types of food abundance. Food is plentiful when there is a lot of it in a given area (left). Food is scarce when there is not very much of it in a given area (right). Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-6\/\">Food abundance and food scarcity (Figure 6.7)<\/a> by Karin Enstam Jaffe 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<figure style=\"width: 690px\" class=\"wp-caption aligncenter\"><img style=\"font-size: 1em\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-2.png\" alt=\"Three squares with dots in different formations. \" width=\"690\" height=\"214\" \/><figcaption class=\"wp-caption-text\">Figure 7.6a-c: Three types of food distribution. a. Food has a uniform distribution when it is spread out evenly in the environment. b. Food has a clumped distribution when it is found in patches. c. Food is randomly distributed when it has neither uniform nor clumped distribution. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-6\/\">Food distribution patterns (Figure 6.8)<\/a> by Karin Enstam Jaffe 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<h3 class=\"import-Normal\"><strong>Community Ecology<\/strong><\/h3>\n<p class=\"import-Normal\">Primates are members of broader ecological communities composed of other species, including other primates, predators, parasites, and even humans. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_966\">Community ecology<\/a><\/strong> deals with the relationships and interactions between different organisms that occupy the same habitat. Interactions with <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_968\">conspecifics<\/a> <\/strong>(members of the same species) and <strong>heterospecifics<\/strong> (members of different species) are critical aspects of ecological communities. Some habitats support highly diverse <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_972\">primate communities<\/a><\/strong> consisting of 10 or more primate species. How can so many primate species occupy the same area and avoid competition? In most cases, the primate species that live together occupy different <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_974\">niches<\/a><\/strong>, which means they do not meet their needs for food and shelter in the exact same way. Two species can avoid competition by eating different kinds of food, living at different levels of a forest, or even searching for food at different times of day. Because tropical rainforests, like Manu National Park in Peru, are highly variable, with many habitats and many sources of food and shelter, there are many different niches for multiple species to exploit, and large primate communities can result (Figure 7.7).<\/p>\n<figure style=\"width: 710px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5-1-1.png\" alt=\"Eight primate species.\" width=\"710\" height=\"550\" \/><figcaption class=\"wp-caption-text\">Figure 7.7: Eight of the 14 primate species in Manu National Park, Peru. Top row, left to right: Goeldi\u2019s marmoset (Callimico goeldi), Rio Tapaj\u00f3s saki (Pithecia irrorata), tufted capuchin (Sapajus apella); middle row, left to right: emperor tamarin (Saguinus imperator), black-headed night monkey (Aotus nigriceps), Bolivian red howler (Alouatta sara); bottom row, left to right: black-capped squirrel monkey (Saimiri boliviensis), Peruvian spider monkey (Ateles chamek). Credit: Primate species in Manu National Park original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Jaffe 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:\/\/www.flickr.com\/photos\/31223088@N08\/5582747190\/\">Tamarin Baby\/Goeldi\u2019s Monkey<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/31223088@N08\/\">stefan_fotos<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\">CC BY 2.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Pithecia_irrorata_-Brazil-8b.jpg\">Pithecia irrorata -Brazil-8b<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/9092428@N04\">Ana_Cotta<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/deed.en\">CC BY 2.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Tufted_capuchin_on_a_branch_in_Singapore.jpg\">Tufted Capuchin on a Branch in Singapor<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Basile_Morin\">Basile_Morin<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Tamarin_portrait.JPG\">Tamarin Portrait<\/a> by <a href=\"https:\/\/sites.google.com\/site\/thebrockeninglory\/?pli=1\">Brocken Inaglory<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/deed.en\">CC BY-SA 3.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Aotus_nigriceps_1.jpg\">Aotus nigriceps 1<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/dusantos_bh\/\">DuSantos<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/deed.en\">CC BY 2.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Alouatta_sara_%28Bolivian_red_howler%29.jpg\">Aloutta sara (Bolivian Red Howler)<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/nacho_dayz\/\">Raul Ignacio<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\">CC BY-SA 2.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Black-capped_squirrel_monkey_%28Chalalan%29.jpg\">Black-Capped Squirrel Monkey (Chalalan)<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Rodrigo_Mariaca\">Rodrigo Mariaca<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0 License<\/a>; <a href=\"https:\/\/www.flickr.com\/photos\/eye1\/3185562151\/\">Maquisapa (Spider Monkey)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/eye1\/\">Ivan Mlinaric<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\">CC BY 2.0 License<\/a>].<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Competitive Interactions<\/em><\/h4>\n<p class=\"import-Normal\">Although species living in the same location often occupy different niches to avoid competition, when a resource that is important for survival or reproduction is scarce, individuals will compete to obtain that resource. This is a central tenet of Charles Darwin\u2019s theory of evolution by natural selection (see Chapter 2 &amp; 3). Competition between primates takes two forms: Individuals engage in <strong>direct competition<\/strong>, which involves physical interaction between individuals (such as fighting), over resources that are large and worth defending (fruit is a good example of a food resource over which primates will fight). Individuals engage in <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_978\">indirect competition<\/a><\/strong>, in which there is no physical interaction between individuals, when a resource is small. Primates often engage in indirect competition for insects, like grasshoppers, that are eaten quickly, often before another individual arrives on the scene. Primates may engage in direct and\/or indirect competition with members of their own group, with members of other groups of conspecifics, or with heterospecifics.<\/p>\n<h4 class=\"import-Normal\"><em>Predator-Prey Interactions<\/em><\/h4>\n<p class=\"import-Normal\">The plants and animals that primates eat are an important part of their ecological community. In addition to insects, many primates incorporate some <strong>vertebrate<\/strong> (animals with an internal spinal column or backbone) prey into their diet. Often, predation by primates is opportunistic, occurring because the prey happens to be in the right place at the right time. I\u2019ve observed vervets (<em>Chlorocebus pygerythrus<\/em>) opportunistically killing lizards by smashing them against a rock or tree trunk and eating them. More rarely, hunting is deliberate and cooperative. In some chimpanzee (<em>Pan troglodytes<\/em>) populations, hunts involve multiple individuals, each of whom plays a specific role and is rewarded afterward with a share of the prey that has been captured (Samuni Et al. 2018).<\/p>\n<p class=\"import-Normal\">All primates are susceptible to predation by mammalian <strong>carnivores <\/strong>(animals whose diet consists primarily of animal tissue (e.g., Figure 7.8a), reptiles (e.g., Figure 7.8b), or birds of prey (e.g., Figure 7.8c). Although the specific predators found in an ecological community differ based on geography, smaller primates always fall prey to a wider range of predators. Because predators are diverse in their hunting tactics, primates have evolved a wide range of tactics to avoid or escape them. We will discuss some of these behavioural adaptations later in this chapter in the section titled \u201cWhy Do Primates Live in Groups?.\u201d<\/p>\n<p>&nbsp;<\/p>\n<\/div>\n<figure id=\"attachment_210-15\" aria-describedby=\"caption-attachment-210-15\" style=\"width: 765px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-195\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.8-scaled-1.jpg\" alt=\"A leopard, python, and harpy eagle.\" width=\"765\" height=\"229\" \/><figcaption id=\"caption-attachment-210-15\" class=\"wp-caption-text\">Figure 7.8a-c: Examples of primate predators: the Indian leopard (Panthera fusca) is an example of a mammalian carnivore (top left), the South African python (Python natalensis) is an example of a reptilian predator (bottom left), and the harpy eagle (Harpia harpyja) of Central and South America is an example of a bird of prey (right). Credit: a. <a href=\"https:\/\/www.flickr.com\/photos\/srikaanth-sekar\/9814267145\/\">Leopard<\/a> by<a href=\"https:\/\/www.flickr.com\/people\/srikaanth-sekar\/\"> Srikaanth Sekar<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/\"> CC BY-SA 2.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Python_natalensis_G._J._Alexander.JPG\">Python natalensis G. J. Alexander<\/a> by Graham J. Alexander, University of the Witwatersrand, USGS, is in the<a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\"> public domain<\/a>. c. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Harpy_Eagle_clutching_captured_bird_-_Itirapina_Reserve.jpg\">Harpy Eagle clutching captured bird - Itirapina Reserve<\/a> by Jonathan Wilkins has been modified (cropped) and 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<div class=\"__UNKNOWN__\">\n<h4 class=\"import-Normal\"><em>Mutualistic Interactions<\/em><\/h4>\n<p class=\"import-Normal\">So far, we've discussed competitive and predator-prey interactions in primate communities. But there are some interactions (between different primate species and between primates and other species) that are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_984\">mutualistic<\/a><\/strong>, which is when organisms of different species work together, each benefiting from the interaction or relationship. One example is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_986\">seed dispersal<\/a><\/strong>, which is the process by which seeds move away from the plant that produced them in preparation for germination and becoming a new plant. When seeds are dispersed by animals, like primates, it is an example of mutualism. The primate eats the fruit of a plant, which provides nutrients for its body, and in the process ingests the plant\u2019s seeds. Later, it deposits the seeds at another location as a pile of fertilizer.<\/p>\n<p class=\"import-Normal\">Another example of mutualism is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_988\">polyspecific associations<\/a><\/strong>, which are associations between two or more different species that are maintained by behavioural changes by at least one of the species. While some associations are short in duration, others are semi-permanent. The mutualistic benefits of polyspecific associations include one species gaining access to food that would otherwise have been inaccessible or being alerted to the presence of predators that they would not have not have known were present otherwise. In some cases, individuals seem to recognize and seek out specific members of another species. Twenty years of observations on chimpanzees and Western lowland gorillas (<em>Gorilla gorilla gorilla<\/em>) in the Republic of Congo has revealed social ties (some might call them friendships) between individual chimpanzees and gorillas that last for years and occur in a variety of social contexts, including play (Sanz Et al. 2022).<\/p>\n<h4 class=\"import-Normal\"><em>Parasite-Host Interactions<\/em><\/h4>\n<p class=\"import-Normal\">Primates are hosts for a variety of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_990\">parasites<\/a><\/strong>, which are organisms that live in or on another organism (the host). Parasites come in many forms and pose varying levels of danger to the host. Blood parasites cause diseases like yellow fever and malaria. Skin parasites include fleas and ticks, which feed on the host\u2019s blood, and botflies, which lay eggs in the host\u2019s flesh. Bot fly larvae feed on the host\u2019s flesh as they develop and eventually (if not removed) break through the skin at maturity. Gut parasites, like tapeworms, get into the intestines and feed off of the food that is being digested by the host. Because most primates live in groups (see the \u201cPrimate Societies\u201d section of this chapter), the tendency for <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_992\">social transmission<\/a><\/strong> of parasites, or the transfer of parasites from one individual to another, is high. Primates have evolved mechanisms to avoid parasite infection, including switching sleeping and feeding sites so as to avoid parasites. Mandrills (<em>Mandrillus sphinx<\/em>) have been shown to avoid grooming infected conspecifics as well as to avoid their feces, which smell different than the feces of individuals who are not infected with parasites (Poirotte Et al. 2017). Other primates, including chimpanzees, appear to self-medicate when infected with parasites by ingesting plants that have antiparasitic properties (Krief Et al. 2005).<\/p>\n<h4 class=\"import-Normal\"><em>Human-Primate Interactions<\/em><\/h4>\n<p class=\"import-Normal\">Humans are part of many primate communities and our relationship with our closest relatives is often complicated. In some areas, humans hunt primates for their meat or as trophies, or so they can sell the infants as pets. As the human population increases in size, our demand for natural resources, like wood to build houses or land on which to grow food, also increases, often at the expense of pristine primate (and other animal) habitat. As their natural habitat shrinks, primates search for food in areas occupied by humans and may be shot as crop-raiding pests. While deforestation, hunting, and the pet trade are examples of ways in which humans negatively affect the lives of primates, some human-primate interactions are beneficial. In some parts of the world primates are central to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_994\">ecotourism<\/a><\/strong>, which focuses on nature-based attractions to educate tourists and uses economically and ecologically sustainable practices. Perhaps one of the greatest success stories of ecotourism involves the mountain gorillas of Rwanda (see Figure 7.4b). After internal conflict plagued Rwanda during the 1990s, the Virunga Mountains area developed gorilla-based tourism to aid in socioeconomic development and to bring stability to the region. This process not only helped to increase mountain gorilla populations but was also able to generate enough income to cover the operation costs of three national parks and provide income and other benefits to people living in the area (Maekawa Et al. 2013). You can learn more about human-primate interactions in Appendix B: Primate Conservation.<\/p>\n<h2 class=\"import-Normal\">Primate Societies<\/h2>\n<p class=\"import-Normal\">Unlike many other animals, primates are highly social and many live in stable groups consisting of adult males and females, even outside the<strong> <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_996\">breeding season<\/a><\/strong>, when females are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_998\">receptive<\/a><\/strong> and available for mating because they are not pregnant or nursing. Indeed, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1000\">sociality<\/a><\/strong>, or the tendency to form social groups, is a key behavioural adaptation of the order primates (see Chapter 6). This has led primatologists to ask two questions: \u201cWhy do primates live in groups?\u201d and \u201cWhat types of groups do primates live in?\u201d<\/p>\n<h3 class=\"import-Normal\"><strong>Why Do Primates Live in Groups?<\/strong><\/h3>\n<p class=\"import-Normal\">Primates live in groups when the benefits of doing so exceed the costs. Although there are many potential benefits to group living, enhanced feeding competition and predator avoidance are important benefits for many group living primates. When primates feed on high-quality, scarce food (like fruit), larger groups are more successful in competition with other groups. For example, in a long-term study of vervets in Kenya\u2019s Amboseli National Park, larger vervet groups had larger and better <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1002\">home ranges<\/a><\/strong>, which is the area in which a group regularly moves around as it performs its daily activities, including searching for food and water. Females in larger groups had higher average infant and female survival rates than the smallest group. Because pregnancy and nursing are energetically expensive for females, female <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1004\">reproductive success<\/a><\/strong>, or genetic contribution to future generations (measured by the number of offspring produced), is limited by access to food. Although living in a group means females compete with members of their own group for food, the benefits of being a member of a larger vervet group outweigh the costs (Cheney &amp; Seyfarth 1987).<\/p>\n<p class=\"import-Normal\">However, because they contain more individuals, larger groups are more likely to attract the attention of predators compared to smaller groups. This is one of the reasons that primates who rely on <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1006\">crypsis<\/a><\/strong>, or the ability to avoid detection by others, including predators, are often <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1008\">solitary<\/a> <\/strong>(the term used to describe individuals who do not live together with other members of their species) and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1010\">nocturnal<\/a><\/strong>, or active at night. If an animal is already hard to see because it is active at night, then moving quietly in small groups is a good strategy to avoid detection by predators. The slow loris (<em>Nycticebus coucang<\/em>) of Southeast Asia is a good example of this strategy (Figure 7.9a). Nocturnal and solitary, the slow loris moves slowly and quietly as its primary strategy to avoid detection (Wiens &amp; Zitzmann 2003). In contrast, primates who live in large groups and are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1012\">diurnal<\/a><\/strong>, or active during the day (like gelada baboons [<em>Theropithecus gelada<\/em>]; Figure 7.9b) cannot avoid detection by predators. Instead, group-living primates rely on behaviours that alert others to the presence of danger and\/or deter predators, including shared <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1014\">vigilance<\/a><\/strong> (watchful behaviour to detect potential danger), <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1016\">mobbing<\/a><\/strong> (the act of cooperatively attacking or harassing a predator), and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1018\">alarm calling<\/a><\/strong> (vocalizations emitted by social animals in response to danger). We will discuss alarm calls in the <em>Communication<\/em> section.<\/p>\n<figure id=\"attachment_196-3\" aria-describedby=\"caption-attachment-196-3\" style=\"width: 761px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-196\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.9.jpg\" alt=\"A slow loris. A group of gelada baboons.\" width=\"761\" height=\"269\" \/><figcaption id=\"caption-attachment-196-3\" class=\"wp-caption-text\">Figure 7.9a\/b: Some primates, like the slow loris (left), are solitary and spend most of their time alone. However, most primates, like the gelada baboon (right), live in groups of varying sizes. Credit: a. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Slow_Loris.jpg\">Slow Loris<\/a> by Jmiksanek is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\"> CC BY-SA 3.0 License<\/a>. b. <a href=\"https:\/\/www.flickr.com\/photos\/39997856@N03\/7588490544\">Field of baboons<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/39997856@N03\/\">mariusz kluzniak<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/2.0\/\">CC BY-NC-ND 2.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>What Types of Groups Do Primates Live In? <\/strong><\/h3>\n<p class=\"import-Normal\">Primates vary with regard to the types of groups in which they live. A <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1020\">social system<\/a> <\/strong>describes a set of social interactions and behaviours that is typical for a species. The components that make up a species\u2019 social system include:<\/p>\n<ul>\n<li class=\"import-Normal\">Group size, which refers to the number of individuals that typically live together. Primate group size can be highly variable, ranging from one or a few individuals, to a few dozen, upward to several hundred individuals.<\/li>\n<li class=\"import-Normal\">Group composition describes group membership in terms of age class (e.g., adult, juvenile, infant) and sex. In some primates, groups consist of a mother and her dependent offspring while in others, one adult male lives long-term with one adult female and their dependent offspring. In other species, one or more adult males live with multiple females and their offspring.<\/li>\n<li class=\"import-Normal\">A species\u2019 <strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1022\">mating system<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"> refers to which male(s) and female(s) mate. The terms that describe a mating system (e.g., <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1024\">polygyny<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">, in which one male mates with multiple females) are sometimes used to describe a primate species\u2019 social system, but a mating system is one component of the species\u2019 social system. For example, two species might both have polygynous mating systems, but in one species, the group is composed of one male and multiple females, while members of the other species live as solitary individuals.<\/span><\/li>\n<li class=\"import-Normal\"><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1026\">Ranging behaviour<\/a> <\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">refers to the way in which animals move about their environment. Most primate species have a home range, where they perform their daily activities. Some primates defend a <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1028\">territory<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"> which is the part of the home range that the group actively guards in an attempt to keep out conspecifics.<\/span><\/li>\n<li class=\"import-Normal\"><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1030\">Dispersal<\/a> <\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">patterns describe which sex moves to a new group to reproduce. In most primate species, males disperse because the benefits of dispersal, including increased access to mates and reduced competition from other males, outweigh the costs of migrating into a new group, which often comes with aggression from current group members. For many female primates, the opposite is true: females usually benefit from remaining <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1032\">philopatric<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">, or in the group of their birth. This allows them to maintain strong alliances with female relatives, which helps them compete successfully against other groups for food. In solitary species, offspring of both sexes leave their mother\u2019s home range and become solitary. If this did not happen, the species would not be solitary. Even though both sexes disperse in solitary species, males usually disperse farther than females.<\/span><\/li>\n<li class=\"import-Normal\">Social interactions describe the ways in which individuals interact with members of their own and other groups of conspecifics. <strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1034\">Affiliative<\/a> <\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">(i.e., friendly or nonaggressive) behaviours include <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1036\">grooming<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"> (picking through the fur of another individual), playing, or <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1038\">coalitions<\/a> <\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">(temporary alliances between individuals). <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1040\">Agonistic<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"> (i.e., aggressive) behaviours include fighting over food or fighting over access to mates. In groups that contain multiple adult individuals of the same sex, it is common to have a <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1042\">dominance hierarchy<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">, or a group of individuals that can be ranked according to their relative amount of power over others in the hierarchy. Initially, dominance hierarchies are established through the outcome of conflicts. Individuals who lose conflicts with others are <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1044\">subordinate<\/a><\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"> (or low rank) to those who win them. Those who win conflicts are <\/span><strong style=\"text-align: initial;text-indent: 18pt;font-size: 1em\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_744\">dominant<\/a> <\/strong><span style=\"text-align: initial;text-indent: 18pt;font-size: 1em\">(or high rank). Dominant individuals gain access to resources, like food or mates, before subordinates. Once a hierarchy is established, agonism decreases because everyone \u201cknows their place.\u201d<\/span><\/li>\n<\/ul>\n<p class=\"import-Normal\">The main types of primate social systems are as follows: solitary; single-male, single-female; single-male, multi-female; multi-male, multi-female; fission-fusion; and multi-male, single-female. These types are discussed below.<\/p>\n<h4 class=\"import-Normal\"><em>Solitary<\/em><\/h4>\n<p class=\"import-Normal\">Recall that the term <em>solitary<\/em> is used to describe species in which individuals do not live or travel together with other members of the same species, except for mothers and unweaned offspring. Males typically occupy a large home range or territory that overlaps the home ranges of multiple females, with whom they mate (Figure 7.10a). Because one male mates with multiple females, the mating system of solitary primates is polygyny. Social interactions between adults are limited but because some males do not get to mate, competition between males is intense. When males compete physically, they benefit from large body size and weaponry. The result is<strong> <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1046\">sexual dimorphism<\/a><\/strong>, when males and females look different from one another. Both males and females disperse, although males move farther from their mother than females. The nocturnal West African potto (<em>Perodicticus <\/em><em>potto<\/em>; Figure 7.10b) is solitary. Bornean orangutans, which are diurnal, are also solitary.<\/p>\n<\/div>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_210-16\" aria-describedby=\"caption-attachment-210-16\" style=\"width: 620px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-197\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.10-01-1.jpg\" alt=\"Grouping pattern description is available in caption. A potto in a tree at night also shown.\" width=\"620\" height=\"272\" \/><figcaption id=\"caption-attachment-210-16\" class=\"wp-caption-text\">Figure 7.10a-b: Illustration of a solitary species\u2019 grouping pattern is shown on the left. Key: square = adult male; dot = adult female; open circle represents the outline of the male\u2019s home range; open oval represents individual female home ranges. The West African potto is a solitary primate (right). <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. Polygyny in a Solitary Social System original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Enstam Jaffe is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>. b. <a href=\"https:\/\/www.flickr.com\/photos\/nikborrow\/31307385633\">West African Potto Perodicticus potto Kakum National Park, Ghana<\/a> by<a href=\"https:\/\/www.flickr.com\/people\/nikborrow\/\"> Nik Barrow<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/2.0\/\"> CC BY-NC 2.0 License<\/a>.<\/figcaption><\/figure>\n<div class=\"__UNKNOWN__\">\n<h4 class=\"import-Normal\"><em>Single-Male, Single-Female<\/em><\/h4>\n<p class=\"import-Normal\">Primate species in which an adult male and adult female live together with their dependent offspring have a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1048\">single-male, single-female<\/a><\/strong> social system, sometimes referred to as a \u201cfamily,\u201d with group sizes between two and five individuals. The adult male and adult female engage in behaviours that strengthen their social relationship, or <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1050\">pair bond<\/a><\/strong>, including mutual grooming and resting together. The pair defend a territory (Figure 7.11a) and keep same-sex individuals away from their mate. The adult male and adult female mate with each other, so the mating system is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1052\">monogamy<\/a><\/strong>, although mating outside the pair bond may occur. Species with monogamous mating systems are usually <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1054\">sexually monomorphic<\/a><\/strong> (males and females look similar) because competition for mates is relaxed since most males are able to obtain a mate. Males are usually confident that they are the father of their mate\u2019s infant, so they help with offspring care by carrying the infant when it is not nursing. Once offspring are sexually mature, both males and females disperse. As with solitary species, males disperse farther from their parents than females. Bolivian Gray titi monkeys (<em>Plecturocebus donacophilus<\/em>) are an example of a species that has a single-male, single-female social system. One of their signature behaviours is tail twining, when two individuals sit with their tails wrapped around each other (Figure 7.11b). This behaviour reinforces the social bond among family members and is especially common between the adult male and female. Gibbons (<em>Hylobates<\/em>) and owl monkeys (<em>Aotus<\/em>) also live in single-male, single-female groups.<\/p>\n<figure id=\"attachment_198-3\" aria-describedby=\"caption-attachment-198-3\" style=\"width: 608px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-198\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.11-01.jpg\" alt=\"Left: Circle contains one dot (female) and one square (male). Right: Two titi monkeys.\" width=\"608\" height=\"357\" \/><figcaption id=\"caption-attachment-198-3\" class=\"wp-caption-text\">Figure 7.11a-b: Illustration of a single-male, single-female grouping pattern is shown on the left. Key: square = adult male; dot = adult female; open circle represents the outline of the group\u2019s territory, which the bonded pair defend against conspecifics. The titi monkey (right) is an example of a primate species with a single-male, single-female social system. Credit: a. Single-Male, Single-Female Social System original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Enstam Jaffe is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Callicebus-brunneus-London-Zoo.jpg\">Two Red Titi Monkeys (Callicebus cupreus) sitting together with their tails intertwined at the London Zoo<\/a> by<a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Stevenj\"> Steven G. Johnson<\/a> has been modified (cropped) and 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<\/div>\n<div class=\"__UNKNOWN__\">\n<h4 class=\"import-Normal\"><em>Single-Male, Multi-Female<\/em><\/h4>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1056\">Single-male, multi-female<\/a><\/strong> groups consist of one adult male living with multiple adult females and their dependent offspring (Figure 7.12a ) . These groups can range from as few as five or ten individuals to as many as 50. Female social relationships are governed by the female dominance hierarchy. Females are usually philopatric and males disperse. Males who are unable to join a group of females may join a bachelor group with other males. Because a single male mates with multiple females, the mating system is polygyny. Species that form single-male, multi-female groups may or may not defend a territory, but the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1058\">resident male<\/a><\/strong>, who lives with a group of females, is aggressive toward other males, who may try to take over the group and become the new resident male. Competition between males to be the resident male of a group is intense, and these species usually display sexual dimorphism, with males being larger than females and possessing large canines. Hanuman langurs (<em>Semnopithecus entellus<\/em>) of India form single-male, multi-female groups (Figure 7.12b). When a new male takes over a group of females and ousts the former resident male, he may commit <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1060\">infanticide<\/a>, <\/strong>or kill the unweaned infants. This is especially likely if the new resident male has not yet mated with any of the females and thus cannot be the infants\u2019 father. This causes the females, who were nursing, to become sexually receptive sooner, increasing the new resident male\u2019s chances of producing offspring (Sharma, Ram, and \u200b\u200bRaipurohit 2010). Gorillas, patas monkeys, and golden snub-nosed monkeys (<em>Rhinopithecus roxellana<\/em>) also live in single-male, multi-female groups.<\/p>\n<p>&nbsp;<\/p>\n<\/div>\n<figure id=\"attachment_210-17\" aria-describedby=\"caption-attachment-210-17\" style=\"width: 749px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-199\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.12.jpg\" alt=\"Left: Circle contains nine dots and one square; outside are three squares. Right: Adult langur and infant.\" width=\"749\" height=\"295\" \/><figcaption id=\"caption-attachment-210-17\" class=\"wp-caption-text\">Figure 7.12a-b: An illustration of the one-male, multi-female grouping pattern is shown on the left. Key: square = adult male; dot = adult female; open circle represents the outline of the group\u2019s home range (or territory). The Hanuman langur (right) is an example of a species with a one-male, multi-female social system. Credit: a. Single-Male, Multi-Female Social System original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Enstam Jaffe is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>. b. <a href=\"https:\/\/www.pexels.com\/photo\/close-up-photo-of-two-gray-langurs-8642891\/\">Close-up of Two Grey Langurs<\/a> by<a href=\"https:\/\/www.pexels.com\/@amitrai10\/\"> Amit Rai<\/a> has been modified (cropped) and is<a href=\"https:\/\/www.pexels.com\/license\/\"> free to use via Pexels<\/a>.<\/figcaption><\/figure>\n<div class=\"__UNKNOWN__\">\n<h4 class=\"import-Normal\"><em>Multi-Male, Multi-Female<\/em><\/h4>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1062\">Multi-male, multi-female<\/a><\/strong> groups consist of multiple adult males living with multiple adult females and their dependent offspring. Although there is more than one adult male, there are more adult females than adult males in the group (Figure 7.13a). Multi-male, multi-female groups can range in size from about ten to as many as 500 individuals. They occupy a home range but may or may not defend a territory. In groups that contain multiple males and multiple females, it is not possible for one male to monopolize all the matings, so the mating system is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1064\">polygamy<\/a><\/strong>, in which multiple males mate with multiple females. However, this does not mean that all males have an equal opportunity to mate with all females. In multi-male, multi-female groups, both males and females form a dominance hierarchy. The male dominance hierarchy determines their access to females for mating in much the same way that a female dominance hierarchy determines a female\u2019s access to food. Because their place in the hierarchy can affect their reproductive success, males compete with each other, but because it is rare for males to be excluded from mating altogether, the level of competition and degree of sexual dimorphism are less extreme than what we see in polygynous species. Usually, females are philopatric and males disperse. Vervet monkeys (Figure 7.13b), ring-tailed lemurs (<em>Lemur catta<\/em>), white-faced capuchins (<em>Cebus capucinus<\/em>), and black-capped squirrel monkeys (<em>Saimiri boliviensis<\/em>) live in multi-male, multi-female groups.<\/p>\n<\/div>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_210-18\" aria-describedby=\"caption-attachment-210-18\" style=\"width: 585px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-200\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.13.jpg\" alt=\"Circle contains twelve dots and three squares. Right: Two vervet monkeys.\" width=\"585\" height=\"267\" \/><figcaption id=\"caption-attachment-210-18\" class=\"wp-caption-text\">Figure 7.13a-b: An illustration of the multi-male, multi-female grouping pattern is shown on the left. Key: square = adult male; dot = adult female; open circle represents the outline of the group\u2019s home range (or territory). Vervet monkeys (right) are an example of a species that lives in multi-male, multi-female groups. Credit: a. Multi-Male, Multi-Female Social System original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Enstam Jaffe is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>. b. <a href=\"https:\/\/www.flickr.com\/photos\/berniedup\/6011902081\/\">Vervet Monkeys (Chlorocebus pygerythrus)<\/a> by<a href=\"https:\/\/www.flickr.com\/people\/65695019@N07\"> Bernard DUPONT<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/deed.en\"> CC BY-SA 2.0 License<\/a>.<\/figcaption><\/figure>\n<div class=\"__UNKNOWN__\">\n<h4 class=\"import-Normal\"><em>Fission-Fusion<\/em><\/h4>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1066\">Fission-fusion<\/a><\/strong> is a fluid social system in which the size and composition of the social group changes, with groups splitting (fission) or merging (fusion) depending on food availability (Pinacho-Guendulain &amp; Ramos-Fern\u00e1ndez 2017). When key resources are scarce, individuals spread out (fission) and move and feed individually or in small subgroups (Figure 7.14a). When key food resources are plentiful, individuals come together (fusion) and individuals travel and feed as a more cohesive group (Figure 7.14a). Fission-fusion social structure is believed to reduce feeding competition when resources are scarce. Because group composition changes over time, species with fission-fusion social systems are referred to as a community. Communities consist of multiple adult males, multiple adult females, and offspring, and group size varies but typically ranges from ten to a few dozen individuals. Females typically disperse and males are philopatric. Thus, community males are related and display unusual forms of cooperation. The mating system associated with fission-fusion is polygamy. Because males are not excluded from mating, competition for mates is relaxed and sexual dimorphism is moderate (males are slightly larger than females). Geoffroy\u2019s spider monkeys (<em>Ateles geoffroyi<\/em>) (Figure 7.14b) and chimpanzees both have fission-fusion social system.<\/p>\n<\/div>\n<figure id=\"attachment_210-19\" aria-describedby=\"caption-attachment-210-19\" style=\"width: 720px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-201\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.14.jpg\" alt=\"Diagrams show fission and fusion. Three spider monkeys.\" width=\"720\" height=\"289\" \/><figcaption id=\"caption-attachment-210-19\" class=\"wp-caption-text\">Figure 7.14a-b: An illustration of the fission-fusion grouping pattern appears on the left. The left illustration represents fission, when females travel and feed independently in individual home ranges within the community boundary. The right illustration represents fusion, when community members form a cohesive group. <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>. Key: square = adult male; dot = adult female; open circle represents the outline of the community boundary. Open ovals represent individual female home ranges when the group fissions. Credits: a. Fission-Fusion Social System original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Enstam Jaffe is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>. b. <a href=\"https:\/\/www.flickr.com\/photos\/berniedup\/49562895393\">Geoffry\u2019s Spider Monkeys (Ateles geoffroyi)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/65695019@N07\">Bernard DUPONT<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/deed.en\"> CC BY-SA 2.0 License<\/a>.<\/figcaption><\/figure>\n<div class=\"__UNKNOWN__\">\n<h4 class=\"import-Normal\"><em>Multi-Male, Single-Female<\/em><\/h4>\n<p class=\"import-Normal\">In <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1068\">multi-male, single-female<\/a> <\/strong>groups, two or more males live with one breeding female, her dependent offspring, and non-breeding females (Figure 7.15a). This type of social system is found in the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1070\">callitrichids<\/a><\/strong>, the primate family that includes marmosets (<em>Callithrix<\/em>; Figure 7.15b) and tamarins (<em>Saguinus<\/em>) of Central and South America. Their groups rarely exceed 15 individuals, and each group actively defends their territory from conspecifics. Although more than one adult female may live in the group, the mating system is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1072\">polyandry<\/a><\/strong> because there is only one breeding female who mates with all of the adult males. This is achieved through <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1074\">reproductive suppression<\/a><\/strong>, which involves the breeding female preventing other females from reproducing through physiological and\/or behavioural means (Digby, Ferrari, &amp; Salzman 2011). This limits the opportunities for other females in the group to become pregnant. Instead, these females, and the males in the group, help raise the breeding female\u2019s offspring. This is referred to as <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1076\">cooperative breeding<\/a><\/strong> and usually takes the form of carrying infants, grooming them, and protecting them from danger (de Oliveira Terceiro &amp; Burkart 2019). Because reproductive opportunities for female tamarins and marmosets are limited, they are very competitive, and females are slightly larger than males, which helps them compete for the breeding spot in a group.<\/p>\n<p>&nbsp;<\/p>\n<\/div>\n<figure id=\"attachment_210-20\" aria-describedby=\"caption-attachment-210-20\" style=\"width: 578px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-202\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.15.jpg\" alt=\"Circle contains two squares, two unmarked dots, and a B dot. Right: Marmosets with twins.\" width=\"578\" height=\"269\" \/><figcaption id=\"caption-attachment-210-20\" class=\"wp-caption-text\">Figure 7.15a-b: An illustration of multi-male, single-female grouping pattern appears on the left. Key: square = adult male; B dot = breeding female; unmarked dot = non-breeding female; open circle represents the outline of the group\u2019s territory, which is defended against conspecifics. The common marmoset (Callithrix jacchus) is an example of a primate species that has this type of social system (right). Credit: a. Multi-Male, Single-Female Social System original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Enstam Jaffe is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\"> CC BY-NC 4.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Family_of_Common_Marmoset_-_REGUA_-_Brazil_MG_9480_(12930855765).jpg\">Family of Common Marmoset - REGUA - Brazil MG 9480 (12930855765)<\/a> by<a href=\"https:\/\/www.flickr.com\/people\/30818542@N04\"> Francesco Veronesi<\/a> has been modified (cropped) 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<div class=\"__UNKNOWN__\">\n<h2 class=\"import-Normal\">Reproductive Strategies<\/h2>\n<p class=\"import-Normal\">Reproductive strategies have evolved to maximize individual reproductive success. These strategies can be divided into those that deal with offspring production and care (parental investment) and those that maximize mating success (sexual selection). Because the reproductive physiology of male and female primates differs, males and females differ with regard to parental investment and sexual selection strategies. Female strategies focus on obtaining the food necessary to sustain a pregnancy and choosing the best male(s) to father offspring. Male strategies focus on gaining access to receptive females.<\/p>\n<h3 class=\"import-Normal\"><strong>Parental Investment<\/strong><\/h3>\n<figure style=\"width: 362px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image9.jpg\" alt=\"Monkey holds baby.\" width=\"362\" height=\"241\" \/><figcaption class=\"wp-caption-text\">Figure 7.16: A female Japanese macaque nursing her infant. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Snow_monkey_baby_milk_time.jpg\">Snow monkey baby milk time<\/a> by Daisuke Tashiro 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>Biologically speaking, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1078\">parental investment<\/a><\/strong> is any time or energy a parent devotes to the current offspring that enhances its survival (and eventual reproductive success) at the expense of the parent\u2019s ability to invest in the next offspring (Trivers 1972). Female primates invest more heavily in offspring than males. Even before conception, females produce energy-containing eggs, and they will be responsible for sustaining a fertilized egg until it implants in the uterus. After that, they invest in pregnancy and lactation (Figure 7.16). Because all of this investment requires a lot of energy, female primates can only produce one offspring (or litter) at a time. A species\u2019 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1080\">interbirth interval<\/a><\/strong>, or the typical length of time between one birth and the next, is determined by the length of time necessary to maximize each offspring\u2019s survival without jeopardizing the female\u2019s ability to produce the greatest number of offspring possible. If a female invests too little (i.e., weans an offspring too early), she may give birth to many offspring, but very few (if any) of them will survive. If she invests too much (i.e., nurses an offspring even after it could be weaned), she ensures the survival of that individual offspring but will not be able to produce very many during her lifetime. To maximize her reproductive success, a female must invest <em>just<\/em> long enough to ensure the greatest number of offspring survive to reproduce. We often think of maternal care as an <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1082\">innate<\/a><\/strong> (or natural), instinctive behaviour. Yet this is not the case. The \u201cSpecial Topic: Is Maternal Behaviour Innate?\u201d dispels the myth that maternal behaviour is solely instinctual and explains how female primates learn to be good mothers.<\/p>\n<h3 class=\"import-Normal\"><strong>Sexual Selection<\/strong><\/h3>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1084\">Sexual selection<\/a><\/strong>, or selection for traits that maximize mating success, comes in two forms. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1086\">Intrasexual selection<\/a><\/strong> is selection for traits that enhance the ability of members of one sex to compete amongst themselves (\u201c<em>intra<\/em>sexual\u201d = within one sex). <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1088\">Intersexual selection<\/a><\/strong> is selection for traits that enhance the ability of one sex to attract the other (\u201c<em>inter<\/em>sexual\u201d = between the sexes).<\/p>\n<p class=\"import-Normal\">Intrasexual selection most often operates on males. In the wild, adult females are either pregnant or lactating for most of their adult lives. So, in a given population, there are usually more males available and willing to mate than there are females. The result? Females are a scarce resource over which males compete. Intrasexual selection favours traits that help a male win fights with other males. In primates, these traits include large body size (Figure 7.17a) and large canines (Figure 7.17b). Because females don\u2019t possess these same traits, males and females of some species look different; that is, they are sexually dimorphic (Figure 7.17a).<\/p>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_207-7\" aria-describedby=\"caption-attachment-207-7\" style=\"width: 691px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-204\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.17.jpg\" alt=\"Male baboon with two females. Adult male baboon.\" width=\"691\" height=\"293\" \/><figcaption id=\"caption-attachment-207-7\" class=\"wp-caption-text\">Figure 7.17a-b: a. Hamadryas baboons (Papio hamadryas) are sexually dimorphic. The male (left) is much bigger than the female (centre) and also has different coloured fur. b. Adult males, like this gelada baboon, also have larger canines than females. Credit: a. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Dierenpark_Emmen_baboon_(2679944324).jpg\">Dierenpark Emmen baboon (2679944324)<\/a> by<a href=\"https:\/\/www.flickr.com\/people\/80538772@N00\"> robin bos<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\"> CC BY 2.0 License.<\/a> b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Olive_Baboon_Papio_anubis_in_Tanzania_3066_Nevit.jpg\">Olive Baboon Papio anubis, Picture Taken in Tanzania<\/a> by Nevit Dilmen has been modified (cropped) and 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\">Intersexual selection also tends to operate on males, selecting traits that make a male more attractive to females. Females, in turn, choose among potential fathers. Because female primates invest more in offspring production and care than males (see the \u201cParental Investment\u201d section, above), it is more costly for them if the offspring dies before maturity or reaches maturity but does not reproduce. Thus, it benefits a female primate to be choosy and try to pick the healthiest male as a mate. Males must display traits that tell a female why she should choose <em>him<\/em>, and not another male, as her mate.<\/p>\n<p class=\"import-Normal\">What traits are female primates looking for? In humans, women may look for a mate who can provide important resources, such as food, paternal care, or protection. This is rare in other primates, though, since most females do not need males to provide resources. More commonly, female primates obtain genetic benefits for their offspring from choosing one male over another. Often the specific criteria by which females select mates is unknown. However, if a female chooses a healthy (as indicated by traits like a plush coat, bright colouration, or large body size) or older male, she may obtain genes for her offspring that code for health or long life. If a male\u2019s rank is determined by competitive ability that has a genetic component, females who choose males who win fights may acquire these genes (and qualities) for their offspring. Females in some species appear to prefer new immigrants, sometimes even \u201csneaking\u201d copulations with males who are not established members of their groups. Such a preference may provide their offspring with novel genes and increase genetic variation (for more about the importance of genetic variation, see Chapter 4). Female choice is often more subtle than male-male competition, so it can be more difficult to study. However, as more research is conducted, we continue to improve our understanding of the ways that female primates exert their choice.<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Is Maternal Behaviour Innate?<\/h2>\n<p>Zoos almost always have nurseries where infants are cared for by zookeepers if their mothers will not care for them (Figure 7.18). These exhibits are among the most popular because the babies are so cute and so much fun to watch. And the caretaking positions in zoo nurseries are often among the most coveted by zoo personnel for the same reasons. But if maternal behaviour is instinctive, why do zoo nurseries even exist? The answer is that in many species, including primates, maternal behaviour is not purely instinctual; it is dependent on <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1090\">social learning<\/a><\/strong> (behaviour learned by observing and imitating others), as well.<\/p>\n<figure style=\"width: 433px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image13-1.png\" alt=\"Newborn orangutan feeding from a bottle.\" width=\"433\" height=\"336\" \/><figcaption class=\"wp-caption-text\">Figure 7.18: Newborn orangutan at Audubon Zoo being bottle-fed. Credit: <a href=\"https:\/\/newsroom.audubonnatureinstitute.org\/critically-endangered-orangutan-gives-birth-at-audubon-zoo\/\">Newborn orangutan born at Audubon Zoo being bottle fed<\/a> (2022) by<a href=\"https:\/\/audubonnatureinstitute.org\/\"> Audubon Nature Institute<\/a> is used by permission.<\/figcaption><\/figure>\n<p>Captive female primates, including gorillas and chimpanzees, who have not had the opportunity to observe their mother or other females care for infants do not know how to care for their own offspring. Although it is preferred that the primate mother care for her own infant, there are cases when she will not and humans must step in to ensure the offspring survives. When hand-rearing by humans is necessary, the infant is returned to the group as soon as possible in the hopes that it will learn species-typical behaviour from its mother and other conspecifics. Observations such as these indicate that maternal behaviour is learned, not innate, and that maternal care is critically important to the social and psychological development of young primates.<\/p>\n<\/div>\n<h2 class=\"import-Normal\">Communication<\/h2>\n<p class=\"import-Normal\">In its most basic form, communication occurs when one individual (the sender) emits a signal that conveys information, which is detected by another individual (the receiver). We have discussed several aspects of primate sociality in this chapter, all of which require the communication of information between individuals. But exactly <em>how<\/em> does a female chimpanzee communicate her sexual availability? <em>How<\/em> does a vervet monkey communicate the approach of a leopard or that a python is nearby? <em>How<\/em> do solitary, nocturnal primates, like the slow loris, communicate information about themselves to conspecifics? Primate communication comes in four forms: vocal, visual, olfactory, and tactile. Species vary in their reliance on each.<\/p>\n<h3 class=\"import-Normal\"><strong>Vocal Communication<\/strong><\/h3>\n<p class=\"import-Normal\">Primates use sound to communicate danger or threats, to claim and maintain a territory, or make contact with other group members. Alarm calls are given in response to predators. In some cases, alarm calls are used to alert members of the group to the presence of a predator so they can take evasive action. In other cases, they are directed at the predator itself, signaling that it has been detected. You can learn more about alarm calls as forms of vocal communication in the highlight box in this chapter entitled \u201cDig Deeper: Alarm Calls: Signals to Friends or Foes?.\u201d<\/p>\n<p class=\"import-Normal\">Loud calls are designed to travel great distances and are used in territorial defence by many primate species including indris (<em>Indri indri<\/em>), orangutans, gibbons, and howler monkeys (<em>Alouatta<\/em>). In dense forest, where visual communication can be difficult, loud calls can be useful in signaling to conspecifics that a group or individual occupies a specific area. Howler monkeys are named for their loud calls, or \u201croars,\u201d which can be heard one kilometre or more away (Sch\u00f6n Ybarra 1986). Howler monkey roars may act to maintain distance between neighbouring groups or keep extragroup males from entering the home range (1986).<\/p>\n<p class=\"import-Normal\">Other vocalizations are intended to communicate with individuals in one\u2019s own group. These include vocalizations given as part of threat displays or dominance interactions, as well as contact calls that provide information about one\u2019s location to other group members. Chacma baboons (<em>Papio ursinus<\/em>) have a rich repertoire of vocalizations for communicating with other group members (Fischer Et al. 2008). Adult males give specific vocalizations during threat displays and physical confrontations. Subordinates \u201cscreech\u201d when retreating from a dominant individual, signaling submission. Since baboons rely on membership in their group for finding food and detecting predators, a baboon separated from his group will vocalize in an attempt to regain contact. Young baboons emit their own contact calls when separated from their mothers.<\/p>\n<h3 class=\"import-Normal\"><strong>Visual Communication<\/strong><\/h3>\n<figure id=\"attachment_207-8\" aria-describedby=\"caption-attachment-207-8\" style=\"width: 493px\" class=\"wp-caption alignleft\"><img class=\"wp-image-206\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.19.jpg\" alt=\"Female baboon with sexual swelling. Male and female baboon.\" width=\"493\" height=\"209\" \/><figcaption id=\"caption-attachment-207-8\" class=\"wp-caption-text\">Figure 7.19a-b: Two female hamadryas baboons. The female on the left has a sexual swelling while the female on the right (in foreground, with infant clinging to her belly) does not. An adult male is behind her. Credit: a. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Sexual_swelling_in_female_Hamadryas_baboon.jpg\">Sexual swelling in female Hamadryas baboon<\/a> by<a href=\"https:\/\/www.flickr.com\/photos\/mamoritai\/\"> Mamoritai<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/legalcode\"> CC BY-SA 2.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Hamadryas_baboon_at_Giza_Zoo_by_Hatem_Moushir_36.JPG\">Hamadryas baboon at Giza Zoo by Hatem Moushir 36<\/a> by<a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Hatem_Moushir\"> Hatem Moushir<\/a> has been modified (cropped) and 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><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1092\">Visual communication<\/a><\/strong>, which involves signals that can be seen, is an important component of nonhuman primate behaviour, alone or in combination with other forms of communication. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1094\">Piloerection<\/a><\/strong>, or raising one\u2019s hair or fur, is used in aggressive interactions to make an individual appear larger than it actually is. Female macaques (<em>Macaca<\/em>), baboons (<em>Papio<\/em>), and chimpanzees, signal sexual receptivity through changes in the size, shape, and, often, colour of their hindquarters, called a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1096\">sexual swelling<\/a><\/strong> (Figure 7.19a). The sexual swelling reaches its maximum size at ovulation. When females are not receptive, either because they are pregnant or are nursing, they do not display a sexual swelling (Figure 7.19b). Thus, the presence or absence of a sexual swelling signals a female\u2019s reproductive state.<\/p>\n<p>Monkeys and apes use diverse facial expressions in visual communication. Showing your teeth in a \u201csmile\u201d sends a signal of friendship in humans. Displaying teeth in this way is a sign of anxiety or fear in primates. That male mandrill you see \u201cyawning\u201d at your local zoo is actually displaying his teeth to signal tension or to threaten a rival (Figure 7.20a). In addition to showing their canines, male gelada baboons use \u201clip flips,\u201d in which the gums and teeth are exposed by flipping the upper lip up over the nostrils (Figure 7.20b), and \u201craised eyelids,\u201d in which the pale eyelids are exposed by pulling the scalp back as threatening gestures (Aich, Moos-Heilen, &amp; Zimmerman 1990). Submissive males respond by fleeing or presenting their hindquarters.<\/p>\n<figure id=\"attachment_207-9\" aria-describedby=\"caption-attachment-207-9\" style=\"width: 597px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-207\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.20-1.jpg\" alt=\"Adult male mandrill and adult male hamadryas baboon yawning.\" width=\"597\" height=\"310\" \/><figcaption id=\"caption-attachment-207-9\" class=\"wp-caption-text\">Figure 7.20a-b: Males use visual displays to communicate with other males. The male mandrill (left) is yawning to display his canines, and the male gelada baboon (right) enhances the yawn by flipping his upper lip back and raising his eyelids. Credit: a. <a href=\"https:\/\/pxhere.com\/en\/photo\/559944\">Mandrill<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/mathiasappel\/with\/24639723420\/\">Mathias Appel<\/a> has been modified (cropped) and designated to the <a href=\"https:\/\/creativecommons.org\/share-your-work\/public-domain\/cc0\/\">public domain (CC0)<\/a>. b.<a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:BabouinGeladaAuReveil.JPG\"> BabouinGeladaAuReveil<\/a> by<a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:BluesyPete\"> BluesyPete<\/a> has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\"> CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<figure id=\"attachment_209-3\" aria-describedby=\"caption-attachment-209-3\" style=\"width: 414px\" class=\"wp-caption alignleft\"><img class=\"wp-image-208\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.22.jpg\" alt=\"A bald uakari. A spider monkey.\" width=\"414\" height=\"154\" \/><figcaption id=\"caption-attachment-209-3\" class=\"wp-caption-text\">Figure 7.22a-b: Many monkey species have colourful faces, including the bald uakari (Cacajao calvus; left) and the white-bellied spider monkey (Ateles belzebuth) (right). Credit: a. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Uakari.jpg\">Uakari<\/a> by Coada dragos has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\"> CC BY-SA 4.0 License<\/a>. 6.22b <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Ateles_belzebuth_(White-bellied_spider_monkey)_2.jpg\">Ateles belzebuth (White-bellied spider monkey) 2<\/a> by<a href=\"https:\/\/www.flickr.com\/photos\/ewas-world\/\"> Ewa<\/a> (username: Ewcek65) has been modified (cropped) and is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\"> CC BY 2.0 License<\/a>.<\/figcaption><\/figure>\n<figure style=\"width: 163px\" class=\"wp-caption alignright\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-3.png\" alt=\"A male mandrill\u2019s face.\" width=\"163\" height=\"246\" \/><figcaption class=\"wp-caption-text\">Figure 7.21: The colourful face of the male mandrill provides information about health and fitness to other mandrills.Credit: <a href=\"https:\/\/www.flickr.com\/photos\/mathiasappel\/24842082402\/\">Mandrill<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/mathiasappel\/with\/24639723420\/\">Mathias Appel<\/a> has been modified (cropped) and designated to the <a href=\"https:\/\/creativecommons.org\/share-your-work\/public-domain\/cc0\/\">public domain (CC0)<\/a>.<\/figcaption><\/figure>\n<p>Primates also communicate through colour. In female and male mandrills, facial colouration provides information about an individual\u2019s health, competitive ability, and reproductive state to conspecifics (Figure 7.21; Setchell &amp;t al. 2008; Setchell, Wickings, &amp; Knapp 2006). Variation in facial colouration among monkeys of Central and South America ranges from very simple (Figure 7.22a) to complex (Figure 7.22b). Species living with larger numbers of other primate species have evolved more complex facial colouration patterns, suggesting that this trait evolved as a form of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1098\">species recognition<\/a><\/strong>, or the ability to differentiate conspecifics from members of other species (Santana, Lynch Alfaro, &amp; Alfaro 2012).<\/p>\n<h3 class=\"import-Normal\"><strong>Olfactory Communication<\/strong><\/h3>\n<p class=\"import-Normal\">All primates use scent to communicate. Females secrete chemicals from their <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1100\">anogenital<\/a> <\/strong>region (the area of the anus and genitals) that provide males with information about their reproductive state. In some species, like macaques and chimpanzees, this olfactory signal is enhanced by a sexual swelling, as discussed above. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1102\">Olfactory communication<\/a><\/strong>, or communicating through scent, is particularly important for monkeys of Central and South America, lemurs, and lorises. Male and female common squirrel monkeys (<em>Saimiri sciureus<\/em>) (Figure 7.23a) engage in \u201curine washing,\u201d in which an individual urinates on its hands and feet and then uses them to spread urine all over its body. Urine washing may be used to mark trails for others to follow, to control body temperature, as part of dominance displays, or to communicate reproductive state (Boinski 1992). During aggressive interactions with other males, male ring-tailed lemurs rub their tails with scent from glands on their wrists and chests. They use their \u201cperfumed\u201d tails in aggressive interactions with other males, who may respond by waiving their own scented tail, with physical aggression, or by fleeing (Jolly 1966). Males also waive their tails, saturated in scent, to attract females (Shirasu et al. 2020). Males use scent glands in their wrists to mark territorial boundaries (Figure 7.23b; Mertl-Millhollen 1988).<\/p>\n<\/div>\n<p>&nbsp;<\/p>\n<figure id=\"attachment_210-21\" aria-describedby=\"caption-attachment-210-21\" style=\"width: 641px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-210\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.23.jpg\" alt=\"A squirrel monkey. A ring-tailed lemur.\" width=\"641\" height=\"332\" \/><figcaption id=\"caption-attachment-210-21\" class=\"wp-caption-text\">Figure 7.23a-b: Some primates, like the common squirrel monkey (left) and the ring-tailed lemur (right), communicate using scent. Credit: a. <a href=\"https:\/\/www.flickr.com\/photos\/rubund\/6337874822\/\">Saimiri sciureus<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/rubund\/\">Ruben Undheim<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/\">CC BY-SA 2.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Lemur_catta_004.jpg\">Lemur catta 004<\/a> by <a href=\"https:\/\/en.wikipedia.org\/wiki\/User:Maky\">Maky<\/a> has been modified (cropped) and 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<div class=\"__UNKNOWN__\">\n<h3 class=\"import-Normal\"><strong>Tactile Communication<\/strong><\/h3>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1104\">Tactile communication<\/a><\/strong>, or communicating through touch, is very important in all primate species. Physical contact is used to comfort and reassure, is part of courtship and mating, and is used to establish dominance and alliances. Grooming is an important and clearly enjoyable form of tactile communication for all primates (Figure 7.24). Not only does grooming serve to clean the skin and fur, removing parasites and debris, but it is an important affiliative behaviour that helps reinforce social bonds, repair relationships, and cement alliances.<\/p>\n<figure style=\"width: 674px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image1-8.png\" alt=\"Four primate species grooming.\" width=\"674\" height=\"478\" \/><figcaption class=\"wp-caption-text\">Figure 7.24: Examples of grooming in Japanese macaques (upper left), tufted capuchins (Sapajus apella) (upper right), gelada baboons (lower left), and black-and-white ruffed lemurs (Varecia variegata; lower right). Credit: Examples of grooming original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Karin Jaffe 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:Yakushima_macaques_grooming_each_other.jpg\">Yakushima macaques grooming each other<\/a> by<a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Grendelkhan\"> Grendelkhan<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Tufted_capuchin_monkeys_grooming_session_III.jpg\">Tufted capuchin monkeys grooming session III<\/a> by Adrian Soldati, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>;<a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Baboons_Wunania_012018.jpg\"> Baboons Wunania 012018<\/a> by Kim Toogood, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>;<a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Black-and-white_ruffed_lemur_03.jpg\"> Black-and-white ruffed lemur 03<\/a> by Mattis2412, <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/deed.en\">public domain (CC0 1.0)<\/a>].<\/figcaption><\/figure>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Dig Deeper: Alarm Calls: Signals to Friends or Foes?<\/h2>\n<p class=\"import-Normal\">Alarm calls are common among group-living primates. They often serve to notify conspecifics of potential danger, as is the case with vervet monkeys. Research has shown that: (1) vervets classify predators based on hunting style; (2) alarm calls convey information to other vervets about that hunting style; and (3) other vervets respond in ways appropriate for evading that type of predator (Seyfarth, Cheney, &amp; Marler 1980a). When a vervet gives a \u201cleopard\u201d alarm call (directed at mammalian carnivores like leopards, Figure 7.25a), monkeys on the ground climb the nearest tree, while monkeys already in trees stay there or climb higher. Since most mammalian carnivores hunt on the ground, getting into, and staying in, a tree is the best option for escape. When the \u201csnake\u201d alarm call is given, vervets stand on their hind legs and look down at the ground (Figure 7.25b). Since snakes are not pursuit predators, locating them quickly so as to avoid them is the best strategy. Lastly, when an \u201ceagle\u201d alarm call is given, vervets look up or run into bushes, both of which are useful responses for avoiding hawks and eagles, which attack from above (Figure 7.25c). Vervets clearly understand the meaning of each type of alarm call, as they respond appropriately even when they do not see the actual predator (Seyfarth, Cheney, &amp; Marler 1980b). Such <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1106\">semantic communication<\/a><\/strong>, which involves the systematic use of signals to refer to objects in the environment, was once believed to be unique to humans. It may be a precursor to the symbolic capacities of human language.<\/p>\n<figure style=\"width: 482px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5-1.jpg\" alt=\"Primate in a tree views a leopard.\" width=\"482\" height=\"344\" \/><figcaption class=\"wp-caption-text\">Figure 7.25a<\/figcaption><\/figure>\n<figure style=\"width: 484px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image2-2.jpg\" alt=\"Primate views snake on the ground.\" width=\"484\" height=\"338\" \/><figcaption class=\"wp-caption-text\">Figure 7.25b<\/figcaption><\/figure>\n<figure style=\"width: 481px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-2.jpg\" alt=\"Primate on the ground sees bird.\" width=\"481\" height=\"517\" \/><figcaption class=\"wp-caption-text\">Figure 7.25c\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 Figure 7.25a-c: Vervet monkeys respond in different ways to alarm calls for each of their three main predators (leopards, snakes, and eagles) which are appropriate to predator hunting strategies. Credit: Vervet Monkey Alarm Calls by Mary Nelson, original to Explorations: An Open Invitation to Biological Anthropology, 2nd edition, 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\">Research on other African monkeys indicates that some species use alarm calls to signal to the predator that it has been detected. Diana monkeys (<em>Cercopithecus diana<\/em>) give alarm calls to leopards (<em>Panthera pardus<\/em>) but not chimpanzees (Zuberb\u00fchler, No\u00eb, &amp; Seyfarth 1997). Because leopards are stealth predators, they rely on the element of surprise to sneak up on their prey (Figure 7.26a). Alarm calling at leopards appears to tell the leopard that it has been seen and therefore its chance of success will be low. Research shows leopards are more likely to stop hunting after an alarm call has been emitted. Unlike leopards, chimpanzees are pursuit predators and may even use alarm calls to locate potential prey (Figure 7.26b). With such a predator, prey are better off remaining as silent as possible so as not to alert the predator to their location (Zuberb\u00fchler Et al. 1999).<\/p>\n<figure id=\"attachment_215-3\" aria-describedby=\"caption-attachment-215-3\" style=\"width: 1907px\" class=\"wp-caption alignnone\"><img class=\"wp-image-215 size-full\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/6.26.jpg\" alt=\"Leopard crouches in grass. Chimpanzee looks up. \" width=\"1907\" height=\"589\" \/><figcaption id=\"caption-attachment-215-3\" class=\"wp-caption-text\">Figure 7.26a-b: Because leopards (left) and chimpanzees (right) hunt differently, Diana monkeys react differently to them. Credit: a. <a href=\"https:\/\/www.flickr.com\/photos\/thimindu\/5842997328\">Crouching Leopard<\/a> by<a href=\"https:\/\/www.flickr.com\/photos\/thimindu\/\"> Thimindu Goonatillake<\/a> is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/\"> CC BY-SA 2.0 License<\/a>. b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Chimpanzee_in_the_wild.jpg\">Chimpanzee in the wild<\/a> by D.G. Kulakov 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\">\n<\/div>\n<h2 class=\"import-Normal\"><span style=\"text-align: initial;font-size: 1em\">The Question of Culture<\/span><\/h2>\n<p class=\"import-Normal\">It may be surprising in a chapter on nonhuman primates to see a discussion of culture. After all, culture is considered by many, including cultural anthropologists, to be a distinguishing characteristic of humans. Indeed, some anthropologists question claims of culture in primates and other animals. Definitions of animal culture focus on specific behaviours that are unique to one population. Anthropological definitions of human culture emphasize shared ideology (e.g., values, morals, beliefs) and symbols, not just behaviour. Using this definition, some cultural anthropologists view primates as lacking culture because of the absence of symbolic life (e.g., religion). However, the longer we study primate groups and populations, the more insight we gain into primate behavioural variation. If we define <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1108\">culture<\/a><\/strong> as the transmission of behaviour from one generation to the next through social learning, then we must view at least some of the behavioural variation we see in primates as forms of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1110\">cultural tradition<\/a><\/strong>, or a distinctive pattern of behaviour shared by multiple individuals in a social group that persists over time (Whiten 2001).<\/p>\n<h3 class=\"import-Normal\"><strong>Chimpanzee Culture<\/strong><\/h3>\n<p class=\"import-Normal\">Due to both their high level of intelligence and the large number of long-term studies on several different populations, chimpanzees provide the best example of cultural tradition in primates. Chimpanzees express cultural variation in multiple behavioural patterns, ranging from population-specific prey preferences and hunting strategies to tool-use techniques and social behaviours. For example, in Tanzania, chimpanzees fish for termites by stripping twigs and then poking the twigs into termite mounds. The termites react to the \u201cinvasion\u201d by attacking the twig. The chimpanzee pulls the twig out, termites attached, and eats them. In Gambia, they use modified twigs to extract honey from holes in trees. In Fongoli, S\u00e9n\u00e9gal, chimpanzees use sticks as \u201cspears\u201d that they stab into tree cavities to hunt for galagos (Figure 7.27). Multiple chimpanzee populations use a \u201chammer and anvil\u201d to crack open nuts, but the specific techniques differ. Because the cultural traditions are so diverse and unique, if a researcher can observe enough of a chimpanzee\u2019s behaviour, it is possible to assign that individual to a specific community, much in the same way a human being can be associated with a specific culture based on his or her behaviour (Whiten 2011).<\/p>\n<figure style=\"width: 800px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image3-4.jpg\" alt=\"Chimpanzees hunting galagos by poking them with a stick. \" width=\"800\" height=\"453\" \/><figcaption class=\"wp-caption-text\">Figure 7.27a-d: Tool-assisted hunting by a chimpanzee at Fongoli, S\u00e9n\u00e9gal. An adult male chimpanzee uses a tree branch with a modified end to (a\u2013c) stab into a cavity within a hollow tree branch that houses a galago. He ultimately captures the galago as (d) his adolescent brother looks on. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Pan_troglodytes,_tool_use_in_Senegal.jpg\">Pan troglodytes, tool use in Senegal<\/a> by<a href=\"https:\/\/royalsocietypublishing.org\/content\/2\/4\/140507\"> J. D. Pruetz, P. Bertolani, K. Boyer Ontl, S. Lindshield, M. Shelley, and E. G. Wessling<\/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\">How do chimpanzee cultures develop, and how does cultural transmission occur? Although we do not know for sure how chimpanzee cultural traditions develop initially, it is possible that different groups invent, either accidentally or deliberately, certain behaviours that other individuals copy. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1112\">Immigration<\/a><\/strong>, or movement of an individual into a new group or community, is an important avenue of cultural transmission in chimpanzees, much as it is between human cultures. Immigrants (typically females) may bring cultural traditions to their new community, which residents observe and learn. Conversely, immigrants may observe and learn a cultural tradition practiced in their new community (Whiten 2011).<\/p>\n<h3 class=\"import-Normal\"><strong>Cultural Transmission in Macaques<\/strong><\/h3>\n<p class=\"import-Normal\">Two monkey species are well-known for behavioural variation that has been called \u201cpre-cultural\u201d by some primatologists: Japanese macaques and tufted capuchins (<em>Sapajus apella<\/em>). The transmission of unique <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1114\">foraging<\/a><\/strong> (the act of searching for food) behaviours through the members of a provisioned group of Japanese macaques on Koshima Island is well known (Matsuzawa 2015). In an effort to keep the monkeys nearby, researchers provided them with piles of sweet potatoes. A juvenile female named Imo spontaneously washed a muddy sweet potato in a stream. This new food-processing technique first spread among other juveniles and then gradually to older individuals. Within 30 years, it had spread across generations, and 46 of 57 monkeys in the group engaged in the behaviour. Another example comes from a group living far to the north, in Shiga-Heights, Nagano Prefecture. Researchers used apples to entice Japanese macaques to the area. Within a few years, monkeys visited the area regularly and were observed playing with the water in the hot springs. Soon, they climbed into the hot springs and learned to immerse themselves to keep warm and reduce stress when not foraging (Figure 7.28; Matsuzawa 2018; Takeshita Et al. 2018; recall also our discussion of hot spring use as an example of analogous traits at the beginning of this chapter). These examples share several characteristics with human culture, including invention or modification of behaviour, transmission of behaviour between individuals, and the persistence of the behaviour across generations (McGrew 1998).<\/p>\n<figure style=\"width: 477px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image6-4.png\" alt=\"Two monkeys in a hot spring.\" width=\"477\" height=\"292\" \/><figcaption class=\"wp-caption-text\">Figure 7.28: Hot spring use by Japanese macaques is a culturally transmitted behaviour. Credit: <a href=\"https:\/\/www.flickr.com\/photos\/peterthoeny\/32160301021\">Oooh, This Feels Sooo Good!<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/peterthoeny\/\">Peter Theony - Quality HD Photography<\/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<h2 class=\"import-Normal\">Summary<\/h2>\n<p class=\"import-Normal\">Primates are socially complex, extremely intelligent, and highly adaptable. In this chapter we discussed aspects of primate ecology, including how body size and characteristics of food affect what primates eat and how primates interact with other species in their environment. We examined why primates live in groups, the types of groups in which they are found, and the reproductive strategies used by males and females to maximize reproductive success. Like other aspects of their behaviour, primate communication is varied and complex, and we discussed how primates communicate using vocal, visual, olfactory, and tactile signals. Finally, we explored the question of culture among nonhuman primates and learned that some species have cultural traditions, distinctive patterns of behaviour shared by multiple individuals in a social group that persist over time. Humans and other primates are similar in many ways. Learning about principles of primate ecology and behaviour can help us better understand our own behaviour and the behaviours of our extinct relatives.<\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Review Questions<\/h2>\n<ul>\n<li class=\"import-Normal\">If anthropology is the study of humans, why do some anthropologists study primates?<\/li>\n<li class=\"import-Normal\">How does a primate\u2019s ecology affect their diet and interactions with other organisms?<\/li>\n<li class=\"import-Normal\">Why do primates live in groups and in what types of groups do they live?<\/li>\n<li class=\"import-Normal\">What is parental investment and sexual selection?<\/li>\n<li class=\"import-Normal\">What are some examples of primate communication?<\/li>\n<li class=\"import-Normal\">What is the evidence for cultural traditions in primates and how do primatologists think cultural transmission occurs in primates?<\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\">Key Terms<\/h2>\n<p class=\"import-Normal\"><strong>Abundance<\/strong>: How much food is available in a given area.<\/p>\n<p class=\"import-Normal\"><strong>Adaptation<\/strong>: A trait with a function.<\/p>\n<p class=\"import-Normal\"><strong>Affiliative<\/strong>: Nonaggressive social interactions and associations between individuals.<\/p>\n<p class=\"import-Normal\"><strong>Agonistic<\/strong>: Conflict; aggressive interactions between individuals.<\/p>\n<p class=\"import-Normal\"><strong>Alarm calling<\/strong>: Vocalizations emitted by social animals in response to danger.<\/p>\n<p class=\"import-Normal\"><strong>Analogy<\/strong>: A similar trait found in different species that arose independently.<\/p>\n<p class=\"import-Normal\"><strong>Anogenital<\/strong>: Relating to the anus and genitals.<\/p>\n<p class=\"import-Normal\"><strong>Breeding season<\/strong>: The time of year when females are receptive to mating.<\/p>\n<p class=\"import-Normal\"><strong>Callitrichids<\/strong>: The primate family that includes marmosets and tamarins.<\/p>\n<p class=\"import-Normal\"><strong>Carnivores<\/strong>: Organisms whose diet consists primarily of animal tissue.<\/p>\n<p class=\"import-Normal\"><strong>Coalition<\/strong>: A temporary alliance between individuals.<\/p>\n<p class=\"import-Normal\"><strong>Community ecology<\/strong>: The branch of ecology that deals with the relationships and interactions between different organisms that occupy the same habitat.<\/p>\n<p class=\"import-Normal\"><strong>Comparison<\/strong>: An examination of the similarities and differences between two things, such as two primate species.<\/p>\n<p class=\"import-Normal\"><strong>Conspecifics<\/strong>: Members of the same species.<\/p>\n<p class=\"import-Normal\"><strong>Cooperative breeding<\/strong>: When individuals other than the mother and father help raise the offspring.<\/p>\n<p class=\"import-Normal\"><strong>Crypsis<\/strong>: The ability to avoid detection by other organisms, such as predators.<\/p>\n<p class=\"import-Normal\"><strong>Cultural tradition<\/strong>: A distinctive pattern of behaviour shared by multiple individuals in a social group, which persists over time and is acquired through social learning.<\/p>\n<p class=\"import-Normal\"><strong>Culture<\/strong>: The transmission of behaviour from one generation to the next through observation and imitation.<\/p>\n<p class=\"import-Normal\"><strong>Decolonize<\/strong>: Understanding and highlighting the theory and research of non-Western individuals and perspectives.<\/p>\n<p class=\"import-Normal\"><strong>Descendant<\/strong>: A species that comes after the ancestor species.<\/p>\n<p class=\"import-Normal\"><strong>Direct competition:<\/strong> Competition that involves physical interaction between individuals, such as fighting.<\/p>\n<p class=\"import-Normal\"><strong>Dispersal<\/strong>: To leave one\u2019s group or area. This may or may not involve joining another group.<\/p>\n<p class=\"import-Normal\"><strong>Distribution<\/strong>: How food is spread out.<\/p>\n<p class=\"import-Normal\"><strong>Diurnal<\/strong>: Active during the day.<\/p>\n<p class=\"import-Normal\"><strong>Dominance hierarchy<\/strong>: The ranked organization of individuals established by the outcome of aggressive-submissive interactions.<\/p>\n<p class=\"import-Normal\"><strong>Dominant<\/strong>: Being of high rank.<\/p>\n<p class=\"import-Normal\"><strong>Ecology<\/strong>: The relationship between organisms and their physical surroundings.<\/p>\n<p class=\"import-Normal\"><strong>Ecotourism<\/strong>: A form of tourism that focuses on nature-based attractions to provide learning opportunities and that uses economically and ecologically sustainable practices.<\/p>\n<p class=\"import-Normal\"><strong>Ethology<\/strong>: The study of animal behaviour.<\/p>\n<p class=\"import-Normal\"><strong>Fission-fusion<\/strong>: Societies in which group composition is flexible, such as chimpanzee and spider monkey societies. Individuals may break up into smaller feeding groups (fission) and combine into larger groups (fusion).<\/p>\n<p class=\"import-Normal\"><strong>Fitness<\/strong>: An individual\u2019s ability to survive and reproduce relative to other members of the same species.<\/p>\n<p class=\"import-Normal\"><strong>Folivores<\/strong>: Organisms whose diet consists primarily of leaves.<\/p>\n<p class=\"import-Normal\"><strong>Foraging<\/strong>: The act of searching for food.<\/p>\n<p class=\"import-Normal\"><strong>Frugivores<\/strong>: Organisms whose diet consists primarily of fruit.<\/p>\n<p class=\"import-Normal\"><strong>Grooming<\/strong>: Picking through the fur of another individual for cleaning or bonding purposes.<\/p>\n<p class=\"import-Normal\"><strong>Heterospecifics<\/strong>: Members of different species.<\/p>\n<p class=\"import-Normal\"><strong>Holism<\/strong>: The idea that the parts of a system interconnect and interact to make up the whole.<\/p>\n<p class=\"import-Normal\"><strong>Home range<\/strong>: The area that a group or individual uses over a given period of time (often over a year).<\/p>\n<p class=\"import-Normal\"><strong>Homology<\/strong>: A similar trait found in different species because it was inherited from a common ancestor.<\/p>\n<p class=\"import-Normal\"><strong>Immigration<\/strong>: Movement of an individual into a new group or community.<\/p>\n<p class=\"import-Normal\"><strong>Indirect competition<\/strong>: Competition that does not involve physical interaction between individuals, such as eating food before another individual arrives at the food site.<\/p>\n<p class=\"import-Normal\"><strong>Infanticide<\/strong>: The killing of infants of one\u2019s own species.<\/p>\n<p class=\"import-Normal\"><strong>Innate<\/strong>: Natural; as in behaviour that comes naturally.<\/p>\n<p class=\"import-Normal\"><strong>Insectivores<\/strong>: Organisms whose diets consist primarily of insects.<\/p>\n<p class=\"import-Normal\"><strong>Interbirth interval<\/strong>: The typical length of time between one birth and the next for a species.<\/p>\n<p class=\"import-Normal\"><strong>Intersexual selection<\/strong>: The selection for traits that enhance the ability of the members of one sex to attract the attention of the other.<\/p>\n<p class=\"import-Normal\"><strong>Intrasexual selection<\/strong>: Selection for traits that enhance the ability of members of one sex to compete amongst themselves.<\/p>\n<p class=\"import-Normal\"><strong>Mating system<\/strong>: A way of describing which male(s) and female(s) mate.<\/p>\n<p class=\"import-Normal\"><strong>Metabolism<\/strong>: The chemical changes that take place in an organism that turn nutrients into energy.<\/p>\n<p class=\"import-Normal\"><strong>Mobbing<\/strong>: Cooperatively attacking or harassing a predator.<\/p>\n<p class=\"import-Normal\"><strong>Monogamy<\/strong>: A mating system in which one male mates with one female.<\/p>\n<p class=\"import-Normal\"><strong>Multi-male, multi-female<\/strong>: A group that consists of multiple adult males, multiple adult females, and their dependent offspring.<\/p>\n<p class=\"import-Normal\"><strong>Multi-male, single-female<\/strong>: A group that consists of two or more adult males, one breeding female, their dependent offspring, and non-breeding females.<\/p>\n<p class=\"import-Normal\"><strong>Mutualistic\/mutualism<\/strong>: When different species work together, with each benefiting from the interaction.<\/p>\n<p class=\"import-Normal\"><strong>Niche<\/strong>: The role of a species in its environment; how it meets its needs for food, shelter, etc.<\/p>\n<p class=\"import-Normal\"><strong>Nocturnal<\/strong>: Active at night.<\/p>\n<p class=\"import-Normal\"><strong>Olfactory communication<\/strong>: Conveying information through scent.<\/p>\n<p class=\"import-Normal\"><strong>Omnivores<\/strong>: Organisms whose diet consists of plant and animal matter.<\/p>\n<p class=\"import-Normal\"><strong>Pair bond<\/strong>: A strong, long-term relationship between two individuals.<\/p>\n<p class=\"import-Normal\"><strong>Parasite<\/strong>: An organism that lives in or on another organism.<\/p>\n<p class=\"import-Normal\"><strong>Parental investment<\/strong>: Any time or energy a parent devotes to the current offspring that enhances its survival (and eventual reproductive success) at the expense of the parent\u2019s ability to invest in the next offspring.<\/p>\n<p class=\"import-Normal\"><strong>Philopatric<\/strong>: Remaining in the group of one\u2019s birth.<\/p>\n<p class=\"import-Normal\"><strong>Piloerection<\/strong>: Raising one\u2019s hair or fur in an effort to look bigger.<\/p>\n<p class=\"import-Normal\"><strong>Polyandry<\/strong>: A mating system in which multiple males mate with a single breeding female.<\/p>\n<p class=\"import-Normal\"><strong>Polygamy<\/strong>: A mating system in which multiple males mate with multiple females.<\/p>\n<p class=\"import-Normal\"><strong>Polygyny<\/strong>: A mating system in which one male mates with multiple females.<\/p>\n<p class=\"import-Normal\"><strong>Polyspecific association<\/strong>: An association between two or more different species that involves behavioural changes in at least one of them to maintain the association.<\/p>\n<p class=\"import-Normal\"><strong>Primate community<\/strong>: All primate species that occur in an area.<\/p>\n<p class=\"import-Normal\"><strong>Primatologist<\/strong>: A scientist who studies primate behaviour and\/or ecology.<\/p>\n<p class=\"import-Normal\"><strong>Primatology<\/strong>: The scientific field that studies primate behaviour and\/or ecology.<\/p>\n<p class=\"import-Normal\"><strong>Ranging behaviour<\/strong>: Refers to the way in which animals move about their environment.<\/p>\n<p class=\"import-Normal\"><strong>Receptive<\/strong>: A term used to describe females who are ready for sexual reproduction (i.e., not pregnant or nursing).<\/p>\n<p class=\"import-Normal\"><strong>Reproductive success<\/strong>: An individual\u2019s genetic contribution to future generations, often measured through the number of offspring produced.<\/p>\n<p class=\"import-Normal\"><strong>Reproductive suppression<\/strong>: The prevention or inhibition of reproduction of healthy adults.<\/p>\n<p class=\"import-Normal\"><strong>Resident male<\/strong>: Term that describes the male who lives with a group of females.<\/p>\n<p class=\"import-Normal\"><strong>Seed dispersal<\/strong>: The process by which seeds move away from the plant that produced them in preparation for germination and becoming a new plant.<\/p>\n<p class=\"import-Normal\"><strong>Semantic communication<\/strong>: The systematic use of signals to refer to objects in the environment.<\/p>\n<p class=\"import-Normal\"><strong>Sexual dimorphism<\/strong>: When males and females of a species have different morphological traits.<\/p>\n<p class=\"import-Normal\"><strong>Sexual selection<\/strong>: The selection for traits that increase mating success. This occurs via intersexual selection and intrasexual selection.<\/p>\n<p class=\"import-Normal\"><strong>Sexual swelling<\/strong>: Area of the hindquarters that change in size, shape, and often colour over the course of a female\u2019s reproductive cycle, reaching maximum size at ovulation. Occurs in many primate species that live in Africa and Asia.<\/p>\n<p class=\"import-Normal\"><strong>Sexually monomorphic<\/strong>: When males and females of a species have similar morphological traits.<\/p>\n<p class=\"import-Normal\"><strong>Single-male, multi-female<\/strong>: A group that consists of one adult male, multiple adult female, and their dependent offspring.<\/p>\n<p class=\"import-Normal\"><strong>Single-male, single-female<\/strong>: A group that consists of one adult male, one adult female, and their dependent offspring.<\/p>\n<p class=\"import-Normal\"><strong>Social learning<\/strong>: The idea that new behaviours can be acquired by observing and imitating others.<\/p>\n<p class=\"import-Normal\"><strong>Social system<\/strong>: A way of describing the typical number of males and females of all age classes that live together.<\/p>\n<p class=\"import-Normal\"><strong>Social transmission<\/strong>: Transfer of something from one individual to another; this can include parasites, information, or cultural traditions.<\/p>\n<p class=\"import-Normal\"><strong>Sociality<\/strong>: The tendency to form social groups.<\/p>\n<p class=\"import-Normal\"><strong>Solitary<\/strong>: Living alone.<\/p>\n<p class=\"import-Normal\"><strong>Species recognition<\/strong>: The ability to differentiate conspecifics from members of other species.<\/p>\n<p class=\"import-Normal\"><strong>Subordinate<\/strong>: Being of low rank.<\/p>\n<p class=\"import-Normal\"><strong>Tactile communication<\/strong>: Conveying information through touch.<\/p>\n<p class=\"import-Normal\"><strong>Territory:<\/strong> A home range whose boundary is defended from intrusion by conspecifics.<\/p>\n<p class=\"import-Normal\"><strong>Vertebrates<\/strong>: The group of animals characterized by an internal spinal column or backbone. This includes fish, amphibians, reptiles, birds, and mammals.<\/p>\n<p class=\"import-Normal\"><strong>Vigilance<\/strong>: Watchful behaviour used to detect potential danger, usually in the form of predators or potential competitors.<\/p>\n<p class=\"import-Normal\"><strong>Visual communication<\/strong>: Conveying information through signals that can be seen.<\/p>\n<p class=\"import-Normal\"><strong>Vocal communication<\/strong>: Conveying information through signals that can be heard.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration<\/h2>\n<p class=\"import-Normal\">Goodall, Jane. 1971. <em>In the Shadow of Man<\/em>. Boston: Houghton Mifflin.<\/p>\n<p class=\"import-Normal\">Rowe, Noel, and Marc Myers, eds. 2016. <em>All the World\u2019s Primates. <\/em>Charleston, RI: Pogonias Press.<\/p>\n<p class=\"import-Normal\">Strier, Karen B. 2017. <em>Primate Behavioural Ecology.<\/em> 5th ed. New York: Routledge.<\/p>\n<p class=\"import-Normal\"><a href=\"https:\/\/pin.primate.wisc.edu\/\">Primate Info Net<\/a> is an information service of the National Primate Research Center at the University of Wisconsin, Madison. It includes Primate Factsheets, primate news and publications, a list of primate-related jobs, and an international directory of primatology, among other information.<\/p>\n<p class=\"import-Normal\"><a href=\"https:\/\/www.primate-sg.org\/\">Primate Specialist Group<\/a> is a collection of scientists and conservationists who work in dozens of African, Asian, and Latin American nations to promote research on primate conservation.<\/p>\n<p class=\"import-Normal\">Short videos of some primate behaviours discussed in this chapter:<\/p>\n<ul>\n<li class=\"import-Normal\">Watch vervet monkeys respond to different types of predators: BBC One. n.d. \u201cVervet Monkey\u2019s Escape Plans - Talk to the Animals: Episode 2 Preview.\u201d Accessed December 16, 2022. <a class=\"rId10\" href=\"https:\/\/www.youtube.com\/watch?v=q8ZG8Dpc8mM\">https:\/\/www.youtube.com\/watch?v=q8ZG8Dpc8mM. <\/a><\/li>\n<li class=\"import-Normal\">Watch male gelada baboons use the lip flip in competition with other males: Smithsonian Channel, June 9, 2017. \u201cWhy These Vegetarian Monkeys Have Sharp Predator Teeth.\u201d Accessed July 25, 2019. <a class=\"rId11\" href=\"https:\/\/www.youtube.com\/watch?time_continue=145&amp;v=aC6iYj_EBjY\">https:\/\/www.youtube.com\/watch?time_continue=145&amp;v=aC6iYj_EBjY<\/a>.<\/li>\n<li class=\"import-Normal\">Watch (and listen to!) howler monkeys \u201croar\u201d: Science News. N.d. \u201cHear a Male Howler Monkey Roar.\u201d Accessed November 21, 2022. <a class=\"rId12\" href=\"https:\/\/www.youtube.com\/watch?v=PYar0dkZ6v8\">https:\/\/www.youtube.com\/watch?v=PYar0dkZ6v8<\/a>.<\/li>\n<li class=\"import-Normal\">Watch Japanese macaques using natural hot springs: National Geographic. N.d. \u201cMeditative Snow Monkeys Hang Out in Hot Springs.\u201d Accessed July 25, 2019. <a class=\"rId13\" href=\"https:\/\/www.youtube.com\/watch?v=Aat9O85ynsI\">https:\/\/www.youtube.com\/watch?v=Aat9O85ynsI<\/a>.<\/li>\n<li class=\"import-Normal\">Watch chimpanzees make and use tools: National Geographic. n.d. \u201cChimps and Tools.\u201d Accessed July 25, 2019. <a class=\"rId14\" href=\"https:\/\/www.youtube.com\/watch?v=o2TBicMRLtA\">https:\/\/www.youtube.com\/watch?v=o2TBicMRLtA<\/a>.<\/li>\n<\/ul>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\">Aich, H., R. Moos-Heilen, and E. Zimmermann. 1990. \u201cVocalizations of Adult Gelada Baboons (<em>Theropithecus gelada<\/em>): Acoustic Structure and Behavioural Context.\u201d <em>Folia Primatologica<\/em> 55 (3\u20134): 109\u2013132.<\/p>\n<p class=\"import-Normal\">Bell, Sarah A. 2017. \u201cGaldikas, Birute.\u201d In <em>The International Encyclopedia of Primatology, Volume A\u2013G<\/em>, edited by Agust\u00edn Fuentes, 445\u2013446. Malden, MA: John Wiley &amp; Sons.<\/p>\n<p class=\"import-Normal\">Boinski, S. 1992. \u201cOlfactory Communication among Costa Rican Squirrel Monkeys: A Field Study.\u201d <em>Folia Primatologica<\/em> 59 (3): 127\u2013136.<\/p>\n<p class=\"import-Normal\">Cheney, D. L., and R. M. Seyfarth. 1987. \u201cThe Influence of Intergroup Competition on the Survival and Reproduction of Female Vervet Monkeys.\u201d <em>Behavioral Ecology and Sociobiology<\/em> 21 (6): 375\u2013386.<\/p>\n<p class=\"import-Normal\">de Oliveira Terceiro, Francisco Edvaldo, and Judith M. Burkart. 2019. \u201cCooperative Breeding.\u201d In <em>Encyclopedia of Animal Cognition and Behavior<\/em>, edited by Jennifer Vonk and Todd Shackelford, 1\u20136. Edinburg, Scotland: Springer Cham.<\/p>\n<p class=\"import-Normal\">Digby, Leslie J., Stephen F. Ferrari, and Wendy Saltzman. 2011. \u201cCallitrichines: The Role of Competition in Cooperatively Breeding Species.\u201d In <em>Primates in Perspective<\/em>, edited by Christina J. Campbell, August\u00cdn Fuentes, Katherine C. MacKinnon, Simon K. Bearder, and Rebecca M. Stumpf, 91\u201310. 2nd edition. New York: Oxford University Press.<\/p>\n<p class=\"import-Normal\">Fischer, Julia, Kurt Hammerschmidt, Dorothy L. Cheney, and Robert M. Seyfarth. 2008. \u201cAcoustic Features of Female Chacma Baboon Barks.\u201d <em>Ethology<\/em> 107 (1): 33\u201354.<\/p>\n<p class=\"import-Normal\">Jolly, Alison. 1966. <em>Lemur Behavior: A Madagascar Field Study<\/em>. Chicago: University of Chicago Press.<\/p>\n<p class=\"import-Normal\">Krief, Sabrina, Claude Marcel Hladik, and Claudie Haxaire. 2005. \u201cEthnomedicinal and Bioactive Properties of Plants Ingested by Wild Chimpanzees in Uganda.\u201d <em>Journal of Ethnopharmacology<\/em> 110 (1\u20133): 1\u201315.<\/p>\n<p class=\"import-Normal\">Maekawa, Mkio, Annette Lanjouw, Eug\u00e8ne Rutagarama, and Doublas Sharp. 2013. \u201cMountain Gorilla Tourism Generating Wealth and Peace in Post-Conflict Rwanda.\u201d <em>Natural Resources Forum<\/em> 37 (2): 127\u2013137.<\/p>\n<p class=\"import-Normal\">Matsuzawa, Tetsuro. 2015. \u201cSweet-Potato Washing Revisited: 50th Anniversary of the <em>Primates<\/em> Article.\u201d <em>Primates<\/em> 56: 285\u2013287.<\/p>\n<p class=\"import-Normal\">Matsuzawa, Tetsuro. 2018. \u201cHot-Spring Bathing of Wild Monkeys in Shiga-Heights: Origin and Propagation of a Cultural Behavior.\u201d <em>Primates<\/em> 59: 209\u2013213.<\/p>\n<p class=\"import-Normal\">McGrew, W. C. 1998. \u201cCulture in Nonhuman Primates?\u201d <em>Annual Review of Anthropology<\/em> 27: 301\u2013328.<\/p>\n<p class=\"import-Normal\">Mertl-Millhollen, Anne S. 1988. \u201cOlfactory Demarcation of Territorial but Not Home Range Boundaries by <em>Lemur catta<\/em>.\u201d <em>Folia Primatologica<\/em> 50 (3\u20134): 175\u2013187.<\/p>\n<p class=\"import-Normal\">Pinacho-Guendulain, B., and G. Ramos-Fern\u00e1ndez. 2017. \u201cInfluence of Fruit Availability on the Fission-Fusion Dynamics of Spider Monkeys (<em>Ateles geoffroyi<\/em>).\u201d <em>International Journal of Primatology<\/em> 38: 466\u2013484.<\/p>\n<p class=\"import-Normal\">Poirotte, Cl\u00e9mence, Fran\u00e7ois Massol, Ana\u00efs Herbert, Eric Willaume, Pacelle M. Bomo, Peter M. Kappeler, and Marie J. E. Charpentier. 2017. \u201cMandrills Use Olfaction to Socially Avoid Parasitized Conspicifics.\u201d <em>Science Advances<\/em> 3 (4): e160172.<\/p>\n<p class=\"import-Normal\">Rodrigues, Michelle. 2019. \u201cIt\u2019s Time to Stop Lionizing Dian Fossey as a Conservation Hero.\u201d <em>Lady Science<\/em> website, September 20. Accessed December 14, 2022. <a class=\"rId15\" href=\"https:\/\/www.ladyscience.com\/ideas\/time-to-stop-lionizing-dian-fossey-conservation\">https:\/\/www.ladyscience.com\/ideas\/time-to-stop-lionizing-dian-fossey-conservation<\/a>.<\/p>\n<p class=\"import-Normal\">Samuni, Liran, Anna Preis, Tobias Deschner, Catherine Crockford, and Roman M. Wittig. 2018. \u201cReward of Labor Coordination and Hunting Success in Wild Chimpanzees.\u201d <em>Communications Biology<\/em> 1: 138.<\/p>\n<p class=\"import-Normal\">Santana, Sharlene E., Jessica Lynch Alfaro, and Michael E. Alfaro. 2012. \u201cAdaptive Evolution of Facial Colour Patterns in Neotropical Primates.\u201d <em>Proceedings of the Royal Society B: Biological Sciences<\/em> 279 (1736): 2204\u20132211.<\/p>\n<p class=\"import-Normal\">Sanz, Crickette M., David Strait, Crepin Eyana Ayina, Jean Marie Massamba, Thierry Fabrice Ebombi, Severin Ndassoba Kialiema, Delon Ngoteni, et al. 2022. \u201cInterspecific Interactions Between Sympatric Apes.\u201d i<em>Science<\/em> 25 (10): 105059.<\/p>\n<p class=\"import-Normal\">Sch\u00f6n Ybarra, M. A. 1986. \u201cLoud Calls of Adult Male Red Howling Monkeys (<em>Alouatta seniculus<\/em>).\u201d <em>Folia Primatologica<\/em> 47 (4): 204\u2013216.<\/p>\n<p class=\"import-Normal\">Setchell, Joanna M., Tessa Smith, E. Jean Wickings, and Leslie A. Knapp. 2008. \u201cSocial Correlates of Testosterone and Ornamentation in Male Mandrills.\u201d <em>Hormones and Behavior<\/em> 54 (3): 365\u2013372.<\/p>\n<p class=\"import-Normal\">Setchell, Joanna M., E. Jean Wickings, and Leslie A. Knapp. 2006. \u201cSignal Content of Red Facial Coloration in Female Mandrills (<em>Mandrillus sphinx<\/em>).\u201d <em>Proceedings of the Royal Society B: Biological Sciences<\/em><a class=\"rId16\" href=\"https:\/\/paperpile.com\/b\/Gb7Zko\/Lxpl\"> 273 (1599): 2395\u20132400.<\/a><\/p>\n<p class=\"import-Normal\">Seyfarth, R. M., D. L. Cheney, and P. Marler. 1980a. \u201cMonkey Responses to Three Different Alarm Calls: Evidence of Predator Classification and Semantic Communication.\u201d <em>Science<\/em> 210 (4471): 801\u2013803.<\/p>\n<p class=\"import-Normal\">Seyfarth, Robert M., Dorothy L. Cheney, and Peter Marler. 1980b. \u201cVervet Monkey Alarm Calls: Semantic Communication in a Free-Ranging Primate.\u201d <em>Animal Behaviour<\/em> 28 (4): 1070\u20131094.<\/p>\n<p class=\"import-Normal\">Sharma, Goutam, Chan Ram, and Lal Singh Rajpurohit. 2010. \u201cA Case Study of Infantcide After Resident Male Replacement in <em>Semnopithecus entellus<\/em> around Jodhpur (India).\u201d <em>Proceeding of the Zoological Society<\/em> 63 (2): 93\u201398.<\/p>\n<p class=\"import-Normal\">Shirasu, Mika, Satomi Ito, Akihiro Itoigawa, Takashi Hayakawa, Kodzue Kinoshita, Isao Munechika, Hiroo Imai, and Kazushige Touhara. 2020. \u201cKey Male Glandular Odorants Attracting Female Ring-Tailed Lemurs.\u201d <em>Current Biology<\/em> 30 (11): 2131\u20132138.<\/p>\n<p class=\"import-Normal\">Stanford, Craig B. 2017. \u201cGoodall, Jane.\u201d In <em>The International Encyclopedia of Primatology, Volume A\u2013G<\/em>, edited by Agust\u00edn Fuentes, 471\u2013472. Malden, MA: John Wiley &amp; Sons.<\/p>\n<p class=\"import-Normal\">Stewart, Kelly. 2017. \u201cFossey, Dian.\u201d In <em>The International Encyclopedia of Primatology, Volume A\u2013G<\/em>, edited by Agust\u00edn Fuentes, 432\u2013433. Malden, MA: John Wiley &amp; Sons.<\/p>\n<p class=\"import-Normal\">Takeshita, Rafaela S.C., Fred B. Bercovitch, Kodzue Kinoshita, and Michael A. Huffman. 2018. \u201cBeneficial Effect of Hot Spring Bathing on Stress Levels in Japanese Macaques.\u201d <em>Primates<\/em> 59 (3): 215\u2013225.<\/p>\n<p class=\"import-Normal\">Trivers, Robert L. 1972. \u201cParental Investment and Sexual Selection.\u201d In <em>Sexual Selection and the Descent of Man, 1871\u20131971<\/em>, edited by Bernard Campbell, 136\u2013179. Chicago: Aldine.<\/p>\n<p class=\"import-Normal\">Whiten, Andrew. 2011. \u201cThe Scope of Culture in Chimpanzees, Humans and Ancestral Apes.\u201d <em>Philosophical Transactions of the Royal Society of London B: Biological Sciences<\/em> 366 (1567): 997\u20131007.<\/p>\n<p class=\"import-Normal\">Wiens, Frank, and Annette Zitzmann. 2003. \u201cSocial Structure of the Solitary Slow Loris <em>Nycticebus coucang<\/em> (Lorisidae).\u201d <em>Journal of Zoology<\/em> 261 (1): 35\u201346.<\/p>\n<p class=\"import-Normal\">Zuberb\u00fchler, Klaus, David Jenny, and Redouan Bshary. 1999. \u201cThe Predator Deterrence Function of Primate Alarm Calls.\u201d <em>Ethology<\/em> 105 (6): 477\u2013490.<\/p>\n<p class=\"import-Normal\">Zuberb\u00fchler, Klaus, Ronald No\u00eb, and Robert M. Seyfarth. 1997. \u201cDiana Monkey Long-Distance Calls: Messages for Conspecifics and Predators.\u201d <em>Animal Behaviour<\/em> 53 (3): 589\u2013604.<\/p>\n<h2 class=\"import-Normal\">Acknowledgements<\/h2>\n<p class=\"import-Normal\">The author is grateful to the editors for the opportunity to contribute to this open-source textbook. She thanks Dr. Stephanie Etting for her encouragement and support during the revision of this chapter. Her suggestions, along with comments made by two anonymous reviewers on an earlier draft of this chapter, improved the final version considerably. Finally, she thanks all the primatologists who came before her, especially her advisor, Lynne A. Isbell, for their tireless efforts to understand the behavior and ecology of the living primates. Without their work, this chapter would not have been possible.<\/p>\n<\/div>\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_1683_1714\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1714\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<div class=\"textbox\">\n<h6>Student contributors for this chapter from Salinda Hess' ANTH 203: <em>Emma Costa, Shima Gahima, Will Lefebvre, Audrey Ch\u00e9kina\u00ebl. <\/em><\/h6>\n<p>Learn more about <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/about-the-authors\/\">authors and editors<\/a>,<a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/student-contributors\/\"> Hess' student contributions<\/a>, and <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/versioning-history\/\">versioning history<\/a>.<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\">Learning Objectives<\/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 analyse, 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2023\/05\/image12.jpg\" alt=\"Woman points to dog and fossil on the ground.\" width=\"206\" height=\"248\" \/><figcaption class=\"wp-caption-text\">Figure 8.1: 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 8.1). 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 8.2).<\/p>\n<figure style=\"width: 358px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image22-1.png\" alt=\"Robed figure near a rock structure.\" width=\"358\" height=\"273\" \/><figcaption class=\"wp-caption-text\">Figure 8.2: 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 8.3). 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image14-1.jpg\" alt=\"Rocky coastline and cliffs.\" width=\"283\" height=\"212\" \/><figcaption class=\"wp-caption-text\">Figure 8.3: 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-metre <em>Plesiosaurus<\/em>, considered by many to be her greatest discovery (Figure 8.4). 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image21.png\" alt=\"Plesiosaurus drawing.\" width=\"247\" height=\"375\" \/><figcaption class=\"wp-caption-text\">Figure 8.4: 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 palaeontology. 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_1683_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_1683_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_1683_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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image30-1.jpg\" alt=\"Fossilized shell.\" width=\"264\" height=\"176\" \/><figcaption class=\"wp-caption-text\">Figure 8.5: Murexsul (Miocene): This fossil was found at the Naval Weapons Centre, 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_1683_832\">fossils<\/a><\/strong>, the mineralized copies of once-living organisms (Figure 8.5). Ultimately, these answers provide insights into human evolution. Pay close attention to ways in which modern biological anthropologists use other disciplines to analyse 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_1683_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_1683_836\">eras<\/a>,<\/strong> and eras are divided into <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_838\">periods<\/a><\/strong>. Finally, periods are divided into <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_846\">epochs<\/a><\/strong> (see Figure 8.6; 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-226 size-full\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 8.6: 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_1683_840\">Holocene<\/a><\/strong>. It began almost twelve thousand years ago (<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_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_1683_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 8.7). 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image1.jpg\" alt=\"Two cylindrical towers emitting white steam.\" width=\"299\" height=\"168\" \/><figcaption class=\"wp-caption-text\">Figure 8.7: 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_1683_850\">taphonomy<\/a><\/strong> comes in (Figure 8.8). Taphonomy 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image25-1.jpg\" alt=\"Coyote skull with bones and fur.\" width=\"261\" height=\"348\" \/><figcaption class=\"wp-caption-text\">Figure 8.8: 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 &amp; Wing 1999, 122\u2013141).<\/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 (2004). Additionally, a fresh burnt equid cranium was discovered which supports the theory of mobile hominid scavenging and fire use at the site (2004).<\/p>\n<p>&nbsp;<\/p>\n<div class=\"textbox\">\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"font-family: 'Cormorant Garamond', serif;font-size: 1.602em;font-weight: bold\">Special Topic: Bog Bodies and Mummies<\/span><\/p>\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<figure style=\"width: 357px\" class=\"wp-caption alignright\"><img src=\"https:\/\/upload.wikimedia.org\/wikipedia\/commons\/thumb\/4\/44\/Tollundmannen.jpg\/250px-Tollundmannen.jpg\" alt=\"File:Tollundmannen.jpg\" width=\"357\" height=\"316\" \/><figcaption class=\"wp-caption-text\">Figure 8.9: The head of the bog body known as the Tollund Man, discovered near Tollund, Silkeborg, Denmark, and dated to approximately 375\u2013174 BCE. Credit: <em data-start=\"303\" data-end=\"318\">Tollundmannen<\/em> by Sven Rosborn 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\">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).<\/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_1683_852\">Fossilization<\/a><\/strong>, therefore, is the preservation of an organism against these natural decay processes (Figure 8.10).<\/p>\n<figure style=\"width: 699px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image20-2.png\" alt=\"Five images depicting fossilization.\" width=\"699\" height=\"345\" \/><figcaption class=\"wp-caption-text\">Figure 8.10: 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).<\/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>anaerobic<\/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). 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_1683_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_1683_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 (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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image2-1.jpg\" alt=\"Petrified wood.\" width=\"259\" height=\"194\" \/><figcaption class=\"wp-caption-text\">Figure 8.11: 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_1683_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 colourful and is generally aesthetically pleasing because all the features of the original tree\u2019s composition are illuminated through mineralization (Figure 8.11). 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_1683_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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image7-1.jpg\" alt=\"Partial hominin skeleton on black background.\" width=\"242\" height=\"583\" \/><figcaption class=\"wp-caption-text\">Figure 8.12: \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_1683_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 8.12). 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\"><strong><em>Asphalt<\/em><\/strong><\/h4>\n<figure style=\"width: 510px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image28.jpg\" alt=\"Asphalt lake with mammoth figurines.\" width=\"510\" height=\"340\" \/><figcaption class=\"wp-caption-text\">Figure 8.13: 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image6-3.jpg\" alt=\"Skull with open jaw and large teeth.\" width=\"206\" height=\"245\" \/><figcaption class=\"wp-caption-text\">Figure 8.14: 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 colour. A famous fossil site from California is La Brea Tar Pits in downtown Los Angeles (Figure 8.13). 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_1683_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 8.14). 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\"><strong><em>Igneous Rock<\/em><\/strong><\/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\"><strong><em>Trace Fossils<\/em><\/strong><\/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_1683_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 8.15). 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5-2.jpg\" alt=\"Uneven rock surface with footprints. \" width=\"399\" height=\"245\" \/><figcaption class=\"wp-caption-text\">Figure 8.15: 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_1683_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_1683_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 8.16). Coprolite, like all fossilized materials, can be <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_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 &amp; O\u2019Dea 2014, 133).<\/p>\n<figure style=\"width: 312px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image19-1-1.jpg\" alt=\"Piece of fossilized poop.\" width=\"312\" height=\"224\" \/><figcaption class=\"wp-caption-text\">Figure 8.16: 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\"><strong><em>Pseudofossils<\/em><\/strong><\/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_1683_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 8.17). 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image29.jpg\" alt=\"Rock with black branching fractal veins.\" width=\"426\" height=\"284\" \/><figcaption class=\"wp-caption-text\">Figure 8.17: 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: \u00a0The Power of Poop<\/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<h3 class=\"import-Normal\"><strong>Walking to the Past<\/strong><\/h3>\n<p class=\"import-Normal\">In 1974, British anthropologist Mary Leakey discovered fossilized animal tracks at Laetoli (Figure 8.18), not far from the important paleoanthropological site at Olduvai Gorge in Tanzania. A few years later, a 27-metre 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image13-1-1.png\" alt=\"Eastern Africa map shows sites within Tanzania.\" width=\"495\" height=\"382\" \/><figcaption class=\"wp-caption-text\">Figure 8.18: 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 &amp; 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> (2016).<\/p>\n<figure style=\"width: 482px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image10.jpg\" alt=\"Map shows Tularosa Basin.\" width=\"482\" height=\"331\" \/><figcaption class=\"wp-caption-text\">Figure 8.19: 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 8.19) 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 8.20), 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image31-1.png\" alt=\"Archaeologists on ground. Excavation with footprints. Closeups of footprints.\" width=\"789\" height=\"594\" \/><figcaption class=\"wp-caption-text\">Figure 8.20: 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. For instance, the Acoma Pueblo people have the word for <em>camel<\/em> in their vocabulary. 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<p class=\"import-Normal\"><span style=\"font-family: 'Cormorant Garamond', serif;font-size: 1.602em;font-weight: bold\">Special Topic: Necropolitics<\/span><\/p>\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 centre 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 Canadian Archaeology Association's <em>Principals of Ethical Conduct<\/em>, 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\"><strong>Dating Methods<\/strong><\/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 analyse 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\"><strong><em>Relative Dating<\/em><\/strong><\/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_1683_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_1683_878\">stratigraphy<\/a> <\/strong>(Figure 8.21). 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image11-1.png\" alt=\"Stratigraphic cross-section with 12 strata.\" width=\"382\" height=\"662\" \/><figcaption class=\"wp-caption-text\">Figure 8.21: 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_1683_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 8.21 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_1683_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_1683_884\">cultural dating<\/a> <\/strong>(Figure 8.22). 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image26-1.png\" alt=\"Ax heads, swords, circlets, and pots by type.\" width=\"364\" height=\"557\" \/><figcaption class=\"wp-caption-text\">Figure 8.22: 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_1683_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 &amp; Ezra-Cohn 1959; Goodrum &amp; 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 &amp; Ezra-Cohn 1959; Goodrum &amp; 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 &amp; Olson 2009; Haddy &amp; 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\"><strong><em>Chronometric Dating<\/em><\/strong><\/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_1683_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 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_890\">Elements<\/a>.<\/strong> Each element consists of an <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_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 8.23).<\/p>\n<figure style=\"width: 285px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image18-1-1.png\" alt=\"Atom labeled with nucleus, proton, neutron, and electron.\" width=\"285\" height=\"285\" \/><figcaption class=\"wp-caption-text\">Figure 8.23: 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_1683_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_1683_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_1683_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_1683_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_1683_902\">radiocarbon dating<\/a> <\/strong>(Figure 8.24). 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 &amp; 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image27.png\" alt=\"Creation of Carbon 14.\" width=\"514\" height=\"658\" \/><figcaption class=\"wp-caption-text\">Figure 8.24: 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\">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. For example, annual tree rings (see discussion of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_904\">dendrochronology<\/a><\/strong> below), <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_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 8.25). 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image17-2.jpg\" alt=\"Radiocarbon date calibration curve. \" width=\"547\" height=\"384\" \/><figcaption class=\"wp-caption-text\">Figure 8.25: 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\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_908\">Potassium-argon (K-Ar) dating<\/a><\/strong> and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_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. 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. 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_1683_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 &amp; 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_1683_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_1683_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 &amp; 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_1683_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 8.26), 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image16-1.png\" alt=\"A tree, cross-section of tree core, and tree-ring timeline.\" width=\"364\" height=\"397\" \/><\/p>\n<figure style=\"width: 384px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image23-1-1.png\" alt=\"Tree rings and dates.\" width=\"384\" height=\"396\" \/><figcaption class=\"wp-caption-text\">Figure 8.26: 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 catalogue 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). 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 &amp; Striker 1987).<\/p>\n<div class=\"textbox\" style=\"background: var(--lightblue)\">\n<p><span style=\"font-family: 'Cormorant Garamond', serif;font-size: 1.602em;font-weight: bold\">Special Topic: New Archaeological Evidence Found in Quebec<\/span><\/p>\n<p>Anticosti Island, located in eastern Canada, has emerged in recent years as a site of exceptional paleontological significance. Containing a remarkably well-preserved stratigraphic record, the island hosts over 1,440 fossil species dating back approximately 445 million years. This makes it one of the most complete and continuous marine fossil archives from the Late Ordovician period; a critical interval in Earth\u2019s history marked by the Late Ordovician Mass Extinction (LOME). As the second most ecologically severe extinction event of the Phanerozoic era, LOME resulted in the loss of nearly 85% of marine species (Bond &amp; Grasby, 2020). While previous research has focused on sedimentary records from various global locations, recent discoveries on Anticosti Island have offered compelling new evidence supporting oceanic anoxia as a primary mechanism driving this mass extinction. Research from the UK Natural Environment Research Council (NERC) describes marine anoxia as a drop in seawater oxygen levels, causing marine animals to asphyxiate, \u201ca potent killer that can account for extinctions in benthic groups and deeper-dwelling graptolites and conodonts\u201d (2020, p. 779). Sea-water pyrite sulphate isotope data and analyzing limestone composition are both useful ways in which scientists have gathered this new information, with prominent research published in the <em>Global and Planetary Change<\/em> journal suggesting a potential global perturbation of sulphur cycling during these times of glaciation (Zhang Et al. 2022). While this research is still in its infancy, it supports NERC\u2019s hypothesis that volcanic activity could have caused the second\u2013and most massive\u2013half of the LOME (Bond &amp; Grasby, 2020, p. 780); a warming of the seawater explaining the marine anoxia identified in the sediments. The 2023 designation of Anticosti Island as a UNESCO World Heritage Site underscores its dual significance as both a site of exceptional paleontological value and a place of deep cultural importance. In a CBC interview with Anticosti mayor H\u00e9l\u00e8ne Boulanger, she attributes this recognition to sustained efforts by the Innu communities of Ekuanitshit and Nutashkuan, who have long emphasized the island\u2019s role as a cultural anchor and a repository of ancestral knowledge (Gagn\u00e9-Coulombe, 2023). Anticosti Island now stands as a critical location for advancing scientific understanding of the Late Ordovician Mass Extinction while simultaneously affirming the vital intersection of Indigenous stewardship and global heritage conservation.<\/p>\n<\/div>\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 analyse <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_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 neighbouring 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 &amp; 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_1683_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 &amp; Spaulding 2017; Spaulding &amp; 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\"><strong><em>Global Temperature Reconstruction<\/em><\/strong><\/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_1683_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 8.27).<\/p>\n<figure style=\"width: 389px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 8.27: 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\"><strong><em>Diet Reconstruction<\/em><\/strong><\/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 analyse 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\"><strong><em>Migration<\/em><\/strong><\/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 analysed, 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image24-2.jpg\" alt=\"Upright boulders of Stonehenge.\" width=\"386\" height=\"289\" \/><figcaption class=\"wp-caption-text\">Figure 8.28: 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 8.28). 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 analysed 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-1-2.png\" alt=\"Sites on Yucatan peninsula.\" width=\"455\" height=\"351\" \/><figcaption class=\"wp-caption-text\">Figure 8.29: 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 8.29 and 7.30) 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, thus assuming 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_1683_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 (2022, 68).<\/p>\n<figure style=\"width: 625px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image32-1.png\" alt=\"Cross-section of the Hoyo Negro cenote.\" width=\"625\" height=\"353\" \/><figcaption class=\"wp-caption-text\">Figure 8.30: 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\">Summary<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">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 analyse 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.<\/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: 617px;height: 861px\">\n<thead>\n<tr style=\"height: 24.25pt\">\n<td class=\"Table1-C\" style=\"padding: 5pt;border: 1pt solid #000000;height: 30px;width: 157.257px\">\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: 249.67px\">\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: 165.625px\">\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: 157.257px\">\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: 249.67px\">\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: 165.625px\">\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: 157.257px\">\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: 249.67px\">\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: 165.625px\">\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: 157.257px\">\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: 249.67px\">\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: 165.625px\">\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: 157.257px\">\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: 249.67px\">\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: 165.625px\">\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: 157.257px\">\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: 249.67px\">\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: 165.625px\">\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: 157.257px\">\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: 249.67px\">\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: 165.625px\">\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: 157.257px\">\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: 249.67px\">\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: 165.625px\">\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: 157.257px\">\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: 249.67px\">\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: 165.625px\">\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: 157.257px\">\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: 249.67px\">\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: 165.625px\">\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: 157.257px\">\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: 249.67px\">\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: 165.625px\">\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: 157.257px\">\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: 249.67px\">\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: 165.625px\">\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: 160.59px\"><\/td>\n<td style=\"height: 15px;width: 253.003px\"><\/td>\n<td style=\"height: 15px;width: 168.958px\"><\/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 analyses 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<\/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>Canadian Archaeology Association , <a href=\"https:\/\/canadianarchaeology.com\/caa\/about\/ethics\/principles-ethical-conduct\">\u201cPrinciples of Archaeological Ethics\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<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>Bond, D., &amp; Grasby, S. (2020). Supplemental material: Late Ordovician mass extinction caused by volcanism, warming, and anoxia, not cooling and glaciation. Geology, 48(8), 777\u2013781. https:\/\/doi.org\/10.1130\/geol.26213s.12221825.<\/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. 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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<p>Zhang, J., Lyons, T. W., Li, C., Fang, X., Chen, Q., Botting, J., &amp; Zhang, Y. (2022). What triggered the late Ordovician mass extinction (Lome)? perspectives from geobiology and biogeochemical modeling. Global and Planetary Change, 216. https:\/\/doi.org\/10.1016\/j.gloplacha.2022.103917.<\/p>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Acknowledgements<\/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>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1683_1172\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1172\"><div tabindex=\"-1\"><p>How much food is available in a given area.<\/p>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1683_1693\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1693\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<div class=\"textbox\">\n<p>Learn more about <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/about-the-authors\/\">authors and editors<\/a>,<a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/student-contributors\/\"> Hess' student contributions<\/a>, and <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/versioning-history\/\">versioning history<\/a>.<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\">Learning Objectives<\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li>Illustrate the troubling history of \u201crace\u201d concepts.<\/li>\n<li>Explain human variation and evolution as the thematic roots of biological anthropology as a discipline.<\/li>\n<li>Critique earlier \u201crace\u201d concepts based on a contemporary understanding of the apportionment of human genetic variation.<\/li>\n<li>Explain how biological variation in humans is distributed clinally and in accordance with both isolation-by-distance and Out-of-Africa models.<\/li>\n<li>Identify phenotypic traits that reflect selective and neutral evolution.<\/li>\n<li>Extend this more-nuanced view of human variation to today\u2019s research, the implications for biomedical studies, applications in forensic anthropology, and other social\/cultural\/political concerns.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p>Humans exhibit biological variation. Humans also have a universal desire to categorize other humans in order to make sense of the world around them. Since the birth of the discipline of <strong>biological anthropology, <\/strong>we have been interested in studying how humans vary biologically and what the sources of this variation are. Before we tackle these big problems, first consider this question: Why <em>should<\/em> we study human variation?<\/p>\n<figure style=\"width: 429px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2023\/06\/image26-3.png\" alt=\"Culturally and biologically diverse humans.\" width=\"429\" height=\"429\" \/><figcaption class=\"wp-caption-text\">Figure 14.1: Humans are culturally diverse (in that cultural differences contribute to a great degree of variation between individuals), but those shown are genetically undiverse. (Top left: Hadzabe members in Tanzania; top right: Inuit family; bottom left: Andean man in Peru; bottom right: English woman.) Credit: <a class=\"rId11\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-12\/\">Humans are diverse (Figure 13.1)<\/a> original to <a class=\"rId12\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Michael Rivera is a collective work under a <a class=\"rId13\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0<\/a> license. [Includes <a class=\"rId14\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Tanzania_-_Hadzabe_hunter_(14533536392).jpg\">Tanzania - Hadzabe hunter (14533536392)<\/a> by <a class=\"rId15\" href=\"https:\/\/www.flickr.com\/people\/67947877@N06\">A_Peach<\/a>, <a class=\"rId16\" href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\">CC BY 2.0<\/a>; <a class=\"rId17\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Inuit-Kleidung_1.jpg\">Inuit-Kleidung 1<\/a> by Ansgar Walk, <a class=\"rId18\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0<\/a>; <a class=\"rId19\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Andean_Man.jpg\">Andean Man<\/a> by <a class=\"rId20\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Cacophony\">Cacophony<\/a>, <a class=\"rId21\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0<\/a>; <a class=\"rId22\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Jane_Goodall_GM.JPG\">Jane Goodall GM<\/a> by <a class=\"rId23\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Floatjon\">Floatjon<\/a>, <a class=\"rId24\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">There are certainly academic reasons for studying human variation. First, it is highly interesting and important to consider the evolution of our species (see Chapters 10-13) and how our biological variation may be similar to (or different from) that of other species (e.g., other primates and apes; see Chapters 6 and 7). Second, anthropologists study modern human variation to understand how different biological traits developed over evolutionary time (see Figure 14.1). Suppose we are able to grasp the evolutionary processes that produce the differences in biology, physiology, body chemistry, behaviour, and culture (<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_804\">human variation<\/a><\/strong>). In that case, we can make more accurate inferences about evolution and adaptation among our hominin ancestors, complementing our study of fossil evidence and the archaeological record. Third, as will be discussed in more detail later on, it is important to consider that biological variation among humans has biomedical, forensic, and sociopolitical implications. For these reasons, the study of human variation and evolution has formed the basis of anthropological inquiry for centuries and continues to be a major source of intrigue and inspiration for scientific research conducted today.<\/p>\n<p class=\"import-Normal\">An even-more-important role of the biological anthropologist is to improve public understanding of human evolution and variation\u2014outside of academic circles. Terms such as <strong>race<\/strong> and <strong>ethnicity<\/strong> are used in everyday conversations and in formal settings within and outside academia. The division of humankind into smaller, discrete categories is a regular occurrence in day-to-day life. This can be seen regularly when governments acquire census data with a heading like \u201cgeographic origin\u201d or \u201cethnicity.\u201d Furthermore, such checkboxes and drop-down lists are commonly seen as part of the identifying information required for surveys and job applications.<\/p>\n<p class=\"import-Normal\">According to professors of anthropology, ethnic studies, and sociology, race is often understood as rooted in biological differences, ranging from such familiar traits as skin colour or eye shape to variations at the genetic level. However, race can also be studied as an \u201cideological construct\u201d that goes beyond biological and genetic bases (Fuentes Et al. 2019), at different times relating to our ethnicities, languages, religious beliefs, and cultural practices. Sometimes people associate racial identity with the concept of socioeconomic status or position, or they link ideas about race to what passport someone has, how long they have been in a country, or how well they have \u201cintegrated\u201d into a population.<\/p>\n<p class=\"import-Normal\">Some of these ideas about ethnicity have huge social and political impacts, and notions of race have been part of the motivation behind various forms of racism and prejudice today, as well as many wars and genocides throughout history. <strong>Racism<\/strong> manifests in many ways\u2014from instances of bullying between kids on school playgrounds to underpaid minorities in the workforce, and from verbal abuse hurled at people of colour to violent physical behaviours against those of a certain race. <strong>Prejudice<\/strong> can be characterized as negative views toward another group based on some perceived characteristic that makes all members of that group worthy of disdain, disrespect, or exclusion (not solely along racial lines but also according to [dis]ability, gender, sexual orientation, or socioeconomic background, for example). According to Shay-Akil McLean (2014), \u201cRacism is not something particular to the United States and race is not the same everywhere in the world. Racial categories serve particular contextual purposes depending on the society they are used in, but generally follow the base logic of the supremacy of one type of human body over all others (ordering these human bodies in a hierarchical fashion).\u201d Choosing which biological or nonbiological features to use when discussing race is always a social process (Omi and Winant 2014). Race concepts have no validity to them unless people continue to use them in their daily lives\u2014and, in the worst cases, to use them to justify racist behaviours and problematic ideas about racial difference or superiority\/inferiority. Recent work in anthropological genetics has revealed the similarities amongst humans on a molecular level and the relatively few differences that exist between populations (Omi and Winant 2014).<\/p>\n<p class=\"import-Normal\">The role of the biological anthropologist becomes crucial in the public sphere, because we may be able to debunk myths surrounding human variation and shed light, for the nonanthropologists around us, on how human variation is actually distributed worldwide (see Figure 14.1). Rooted in scientific observations, our work can help nonanthropologists recognize how common ideas about \u201crace\u201d often have no biological or genetic basis. Many anthropologists work hard to educate students on the history of where race concepts come from, why and how they last in public consciousness, and how we become more conscious of racial issues and the need to fight against racism in our societies. Throughout this chapter, I will highlight how humans cannot actually be divided into discrete \u201craces,\u201d because most traits vary on a continuous basis and human biology is, in fact, very <strong>homogenous<\/strong> compared to the greater genetic variation we observe in closely related species. Molecular anthropology, or anthropological genetics, continues to add new layers to our understanding of human biological variation and the evolutionary processes that gave rise to the contemporary patterns of human variation. The study of human variation has not always been unbiased, and thinkers and scientists have always worked in their particular sociohistorical context. For this reason, this chapter opens with a brief overview of race concepts throughout history, many of which relied on unethical and unscientific notions about different human groups.<\/p>\n<\/div>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: My Experiences as an Asian Academic<\/h2>\n<figure style=\"width: 578px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image10-4.jpg\" alt=\"Outdoor photo of this chapter\u2019s author.\" width=\"578\" height=\"433\" \/><figcaption class=\"wp-caption-text\">Figure 14.2: Michael B. C. Rivera in Hong Kong. Credit: Michael B. C. Rivera in Hong Kong original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) is under a <a class=\"rId26\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p>My name is Michael, and I am a biological anthropologist and archaeologist (Figure 14.2). What strikes me as most interesting to investigate is human biological variation today and the study of past human evolution. For instance, some of my research on ancient coastal populations has revealed positive effects of coastal living on dietary health and many unique adaptations in bones and teeth when living near rivers and beaches. I love talking to students and nonscientists about bioanthropologists\u2019 work, through teaching, science communication events, and writing book chapters like this one. I grew up in Hong Kong, a city in southern China. My father is from the Philippines and my mother is from Hong Kong, which makes me a mixed Filipino-Chinese academic. Growing up, I noticed that people came in all shapes, sizes, and colours. My life is very different now in that I have gained the expertise to explain those differences, and I feel a great responsibility to guide those new to anthropology toward their own understandings of diversity.<\/p>\n<p class=\"import-Normal\">Biological anthropology is not taught extensively in Hong Kong, so I moved to the United Kingdom to earn my bachelor\u2019s, master\u2019s, and doctorate degrees. It was fascinating to me that we could answer important questions about human variation and history using scientific methods. However, I did not have many minority academic role models to look up to while I was at university. My department was made up almost exclusively of white westerner faculty, and it was hard to imagine I could one day get a job at these western institutions. While pursuing my degrees, I also remember several instances of my research contributions being overlooked or dismissed. Sometimes professors and fellow students would make racist comments toward Asian scholars (including me) and other Black, Indigenous and researchers of colour, making us greatly uncomfortable in those spaces. When one of us would work up the courage to tell university leaders our experiences of being stereotyped, dismissed, or insulted, we received little support and were further excluded from research and teaching activities. This is a common experience for Black, Indigenous, and other people of colour who pursue biological anthropology, and we face the difficult choice between leaving the field or bearing with such unsafe spaces.<\/p>\n<p class=\"import-Normal\">It became important to me at that time to find other academics of colour with whom to share experiences and form community. I feel inspired by all my colleagues who advocate for greater representation and diversity in universities (whether they are minority academics or not). I admire many of my fellow researchers who are underrepresented and do a great job of representing minority groups through their cutting-edge research and quality teaching at the undergraduate and graduate levels. Although I no longer work in the West, it has remained my great hope that those in the West and the \u201cGlobal North\u201d will continue to improve university culture, and I support any efforts there to welcome all scholars.<\/p>\n<p class=\"import-Normal\">My current work is based in Hong Kong, where I am deeply dedicated to helping develop biological anthropology in East and Southeast Asia and promoting research from our home regions on the international scene. The study of anthropology really highlights how we share a common humanity and history. Being somebody who is \u201cmixed race\u201d and Asian likely played a key role in steering me toward the study of human variation. As this chapter hopefully shows, there is a lot to discuss about race and ethnicity regarding the discipline\u2019s history and current understandings of <strong>human diversity<\/strong>. Some scientific and technological advancements today are misused for reasons to do with money, politics, or the continuation of antiquated ideas. It is my belief, alongside many of my friends and fellow anthropologists, that science should be more about empathy toward all members of our species and contributing to the intellectual and technological nourishment of society. After speaking to many members of the public, as well as my own undergraduate students, I have received lovely messages from other individuals of colour expressing thanks and appreciation for my presence and understanding as a minority scientist and mentor figure. Anthropology needs much more diversity as well as to make room for those who have traveled different routes into the discipline. All paths taken into anthropology are valid and valuable. I would encourage everyone to study anthropology\u2014it is a field for understanding and celebrating the intricacies of human diversity.<\/p>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<h2 class=\"import-Normal\">The History of \"Race\" Concepts<\/h2>\n<h3 class=\"import-Normal\"><strong>\u201cRace\u201d in the Classical Era<\/strong><\/h3>\n<figure style=\"width: 435px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image11-8.png\" alt=\"Painting of four individuals with varied skin colors, head hair, facial hair, and clothing styles.\" width=\"435\" height=\"331\" \/><figcaption class=\"wp-caption-text\">Figure 14.3: (From left to right) Depicting a Berber (Libyan), a Nubian, an Asiatic (Levantine), and an Egyptian, copied from a mural on the tomb of Seti I. Credit: Egyptian races drawing by Heinrich von Minutoli (1820) of a mural by an unknown artist from the tomb of Seti I is in the public domain.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The earliest classification systems used to understand human variation are evidenced by ancient manuscripts, scrolls, and stone tablets recovered through archaeological, historical, and literary research. The Ancient Egyptians had the <em>Book of Gates<\/em>, dated to the New Kingdom between 1550 B.C.E. and 1077 B.C.E (Figure 14.3). In one part of this tome dedicated to depictions of the underworld, scribes used pictures and hieroglyphics to illustrate a division of Egyptian people into the four categories known to them at the time: the Aamu (Asiatics), the Nehesu (Nubians), the Reth (Egyptians), and the Themehu (Libyans). Though not rooted in any scientific basis like our current understandings of human variation today, the Ancient Egyptians believed that each of these groups were made of a distinctive category of people, distinguishable by their skin colour, place of origin, and even behavioural traits.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Another well-known early document is the Bible, where it is written that all humankind descends from one of three sons of Noah: Shem (the ancestor to all olive-skinned Asians), Japheth (the ancestor to pale-skinned Europeans), and Ham (the ancestor to darker-skinned Africans). Similar to the Ancient Egyptians, these distinctions were based on behavioural traits and skin colour. More recent work in historiography and linguistics suggest that the branches of \u201cHamites,\u201d \u201cJaphethites,\u201d and \u201cShemites\u201d may also relate to the formation of three independent language groups some time between 1000 and 3000 B.C.E. With the continued proliferation of Christianity, this concept of approximately three racial groupings lasted until the Middle Ages and spread as far across Eurasia as crusaders and missionaries ventured at the time.<\/p>\n<figure style=\"width: 309px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image9-9.png\" alt=\"Rows of organisms, with plants and animals at the bottom and humans, angels, and God at top.\" width=\"309\" height=\"449\" \/><figcaption class=\"wp-caption-text\">Figure 14.4: The Great Chain of Being from the Rhetorica Christiana by Fray Diego de Valades (1579). Credit: Great Chain of Being 2 by Didacus Valades (Diego Valades 1579) and photographed by Rhetorica Christiana (via Getty Research) is in the public domain.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">There is also the \u201cGreat Chain of Being,\u201d conceived by ancient Greek philosophers like Plato (427\u2012348 B.C.E.) and Aristotle (384\u2012322 B.C.E.). They played a key role in laying the foundations of empirical science, whereby observations of everything from animals to humans were noted with the aim of creating taxonomic categories. Aristotle describes the Great Chain of Being as a ladder along which all objects, plants, animals, humans, and celestial bodies can be mapped in an overall hierarchy (in the order of existential importance, with humans placed near the top, just beneath divine beings; see Figure 14.4). When he writes about humans, Aristotle expressed the belief that certain people are inherently (or genetically) more instinctive rulers, while others are more natural fits for the life of a worker or enslaved person. Based on research by biological anthropologists, we currently recognize that these early systems of classification and hierarchization are unhelpful in studying human biological variation. Both behavioural traits and physical traits are coded for by multiple genes each, and how we exhibit those traits based on our genetics can vary significantly even between individuals of the same population.<\/p>\n<h3 class=\"import-Normal\"><strong>\u201cRace\u201d during the Scientific Revolution<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The 1400s to 1600s saw the beginnings of the <strong>Scientific Revolution<\/strong> in Western societies, with thinkers like Nicholas Copernicus, Galileo Galilei, and Leonardo Da Vinci publishing some of their most important findings. While by no means the first or only scholars globally to use observation and experimentation to understand the world around them, early scientists living at the end of the medieval period in Europe increasingly employed more experimentation, quantification, and rational thought in their work. This is the main difference between the work of the ancient Egyptians, Romans, and Greeks and that of later scientists such as Isaac Newton and Carl Linnaeus in the 1600s and 1700s.<\/p>\n<figure style=\"width: 215px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image24-2.png\" alt=\"Historic painting of a man in 18th-century wig and garments.\" width=\"215\" height=\"259\" \/><figcaption class=\"wp-caption-text\">Figure 14.5: Carl Linnaeus. Credit: <a class=\"rId35\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Carl_von_Linn%C3%A9.png\">Carl von Linn\u00e9<\/a> by <a class=\"rId36\" href=\"https:\/\/en.wikipedia.org\/wiki\/en:Alexander_Roslin\">Alexander Roslin<\/a> (1718-1793) is in the <a class=\"rId37\" 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\">Linnaeus is the author of <em>Systema Naturae<\/em> (1758), in which he classified all plants and animals under the formalized naming system known as <strong>binomial nomenclature<\/strong> (Figure 14.5). This system is <strong>typological<\/strong>, in that organisms are placed into groups according to how they are similar or different to others under study. What was most anthropologically notable about Linnaeus\u2019s typological system was that he was one of the first to group humans with apes and monkeys, based on the anatomical similarities between humans and nonhuman primates. However, Linnaeus viewed the world in line with <strong>essentialism<\/strong>, a problematic concept that dictates that there are a unique set of characteristics that organisms of a specific kind <em>must<\/em> have and that would remove organisms from taxonomic categorizations if they lacked any of the required criteria.<\/p>\n<p>Linnaeus subdivided the human species into four varieties, with overtly racist categories based on skin colour and \u201cinherent\u201d behaviours. Some European scientists during this period were not aware of their own biases skewing their interpretations of biological variation, while others deliberately worked to shape public perceptions of human variation in ways that established \u201c<strong>otherness<\/strong>\u201d and enforced European domination and the subordination of non-European people. The conclusions and claims at which they arrived, consciously or subconsciously, often fit the times they were living through\u2014the so-called <strong>Age of Discovery<\/strong>, when the superiority of European cultures over others was a pervasive idea throughout people\u2019s social and political lives. Although much of Eurasia was linked by spice- and silk-trading routes, the European colonial period between the 1400s and 1700s was marked by many new and unfortunately violent encounters overseas (Figure 14.6). When Europeans arrived by ship on the shores of continents that were already inhabited, it was their first meeting with the Indigenous peoples of the Americas and Australasia, who looked, spoke, and behaved differently from peoples with whom they were familiar. Building on the idea of species and \u201csubspecies,\u201d natural historians of this time invented the term <em>race<\/em>, from the French <em>rasse<\/em> meaning \u201clocal strain.\u201d<\/p>\n<figure style=\"width: 421px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image25-4.png\" alt=\"Spanish explorers with their military and religious gear surrounding indigenous people.\" width=\"421\" height=\"273\" \/><figcaption class=\"wp-caption-text\">Figure 14.6: A painting depicting the colonization of the Mississippi River environs by Spaniard Hernando DeSoto in 1541 (painted in 1853 by William H. Powell). Credit: <a class=\"rId39\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Discovery_of_the_Mississippi.jpg\">Discovery of the Mississippi<\/a> by <a class=\"rId40\" href=\"https:\/\/en.wikipedia.org\/wiki\/en:William_Henry_Powell\">William Henry Powell<\/a> (photograph courtesy of <a class=\"rId41\" href=\"https:\/\/en.wikipedia.org\/wiki\/Architect_of_the_Capitol\">Architect of the Capitol<\/a>) is in the <a class=\"rId42\" href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Another scientist of the times, Johann Friedrich Blumenbach (1752\u20121840), classified humans into five races based on his observations of cranial form variation as well as skin colour. He thus dubbed the \u201coriginal\u201d form of the human cranium the \u201cCaucasian\u201d form, with the idea that the ideal climate conditions for early humans would have been in the Caucasus region near the Caspian Sea. The key insight Blumenbach presented was that human variation in any particular trait should be more accurately viewed as falling along a gradation (Figure 14.7). While some of his theories were correct according to what we observe today with more knowledge in genetics, they erroneously believed that human \u201csubspecies\u201d were \u201cdegenerated\u201d or \u201ctransformed\u201d varieties of an ancestral Caucasian or European race. According to them, the Caucasian cranial dimensions were the least changed over human evolutionary time, while the other skull forms represented geographic variants of this \u201coriginal.\u201d As will be discussed in greater detail later in this chapter, we have genetic and craniometric evidence for sub-Saharan Africa being the origin of the human species instead (see Chapter 13 on the fossil record that places the origins of modern <em>Homo sapiens<\/em> in north and east Africa). Based on work that shows how most biological characteristics are coded for by nonassociated genes, it is not reasonable to draw links between individuals\u2019 personalities and their skull shapes.<\/p>\n<figure style=\"width: 686px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image7-10.png\" alt=\"Historic drawing of five skulls.\" width=\"686\" height=\"259\" \/><figcaption class=\"wp-caption-text\">Figure 14.7: Five skull drawings representing specimens for Blumenbach\u2019s \u201cMongolian,\u201d \u201cAmerican,\u201d \u201cCaucasian,\u201d \u201cMalayan,\u201d and \u201cAethiopian\u201d races. Credit: <a class=\"rId44\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Blumenbach's_five_races.JPG\">Blumenbach's five races<\/a> by Johann Friedrich Blumenbach is in the <a class=\"rId45\" href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>. Original in the 1795 Treatise on \"De generis humani varietate nativa,\" unnumbered page at the end of the book titled \"Tab II\".<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>\u201cRace\u201d and the Dawn of Scientific Racism<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Between the 1800s and mid-1900s, and contrary to what you might expect, an increased use of scientific methods to justify racial schemes developed in scholarship. Differing from earlier views, which saw all humans as environmentally deviated from one \u201coriginal\u201d humankind, classification systems after 1800 became more <strong>polygenetic<\/strong> (viewing all people as having separate origins) rather than <strong>monogenetic<\/strong> (viewing all people as having a single origin). Instead of moving closer to our modern-day understandings of human variation, there was increased support for the notion that each race was created separately and with different attributes (intelligence, temperament, and appearance).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The 1800s were an important precursor to modern biological anthropology as we know it, given that this was when the scientific measurement of human physical features (anthropometry) truly became popularized. However, empirical studies in the 1800s pushed even further the idea that Europeans were culturally and biologically superior to others. While considered one of the pioneers of American \u201cphysical\u201d anthropology, Samuel George Morton (1799\u20121851) was a scholar who had a large role in perpetuating 1800s scientific racism. By measuring cranial size and shape, he calculated that \u201cCaucasians,\u201d on average, have greater cranial volumes than other groups, such as the Indigenous peoples of the Americas and peoples Morton referred to collectively as \u201cNegros.\u201d Today, we know that cranial size variation depends on such factors as Allen\u2019s and Bergmann\u2019s rules, which give a more likely explanation: in colder environments, it is advantageous for those living there to have larger and rounder heads because they conserve heat more effectively than more slender heads (Beals Et al. 1984). The leading figures in craniometry during the 1800s instead were linked heavily with powerful individuals and wealthy sociopolitical institutions and financial bodies. Theories in support of hierarchical racial schemes using \u201cscientific\u201d bases certainly helped continue the exploitative and unethical trafficking and enslavement of Africans between the 1500s and 1800s.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Morton went on to write in his publication <em>Crania Americana<\/em> (1839) a number of views that fit with a concept called <strong>biological determinism<\/strong>. The idea behind biological determinism is that an association exists between people\u2019s physical characteristics and their behaviour, intelligence, ability, values, and morals. If the idea is that some groups of people are essentially superior to others in cognitive ability and temperament, then it is easier to justify the unequal treatment of certain groups based on outward appearances. Another such problematic thinker was Paul Broca (1824\u20121880), after whom a region of the frontal lobe related to language use is named (Broca\u2019s area). Influenced by Morton, Broca likewise claimed that internal skull capacities could be linked with skin colour and cognitive ability. He went on to justify the European colonization of other global territories by purporting it was noble for a biologically more \u201ccivilized\u201d population to improve the \u201chumanity\u201d of more \u201cbarbaric\u201d populations. Today, these theories of Morton, Broca, and others like them are known to have no scientific basis. If we could speak with them today, they would likely try to emphasize that their conclusions were based on empirical evidence and not <em>a priori<\/em> reasoning. However, we now can clearly see that their reasoning was biased and affected by prevailing societal views at the time.<\/p>\n<h3 class=\"import-Normal\"><strong>\u201cRace\u201d and the Beginnings of Physical Anthropology<\/strong><\/h3>\n<p class=\"import-Normal\">In the early 20th century, we saw a number of new figures coming into the science of human variation and shifting the theoretical focus within. Most notably, these included Ale\u0161 Hrdli\u010dka and Franz Boas.<\/p>\n<figure style=\"width: 430px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image19-4.jpg\" alt=\"Historic photo of a middle-aged person in suit and bowtie.\" width=\"430\" height=\"558\" \/><figcaption class=\"wp-caption-text\">Figure 14.8: Ale\u0161 Hrdli\u010dka (1869\u00ad\u20121943), a Czech anthropologist who founded the American Journal of Physical Anthropology. Credit: <a class=\"rId47\" href=\"https:\/\/siarchives.si.edu\/collections\/siris_sic_10822?back=%2Fcollections%2Fsearch%3Fquery%3Dczech%26online%3Dtrue%26page%3D1%26perpage%3D10%26sort%3Drelevancy%26view%3Dlist#\">(Ales Hrdlicka) SIA2009-4246<\/a> (1903) by an unknown photographer <a class=\"rId48\" href=\"https:\/\/www.si.edu\/termsofuse\">is used for educational and non-commercial purposes as outlined by the Smithsonian.<\/a>. [<a class=\"rId49\" href=\"https:\/\/www.si.edu\/\">Smithsonian Institution<\/a> Archives, Record Unit 9521, Box 1, T. Dale Stewart Oral History Interview; and Record Unit 9528, Box 1, Henry B. Collins Oral History Interview.]<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ale\u0161 Hrdli\u010dka (1869\u00ad\u20121943) was a Czech anthropologist who moved to the United States. In 1903, he established the physical anthropology section of the National Museum of Natural History (Figure 14.8). In 1918, he founded the <em>American Journal of Physical Anthropology<\/em>, which remains one of the foremost scientific journals disseminating bioanthropological research. As part of his work and the scope of the journal, he differentiated \u201c<strong>physical anthropology<\/strong>\u201d from other kinds of anthropology: he wrote that physical anthropology is \u201cthe study of racial anatomy, physiology, and pathology\u201d and \u201cthe study of man\u2019s variation\u201d (Hrdli\u010dka 1918). In some ways, although the scope and technological capabilities of biological anthropologists have changed significantly, Hrdli\u010dka established an area of inquiry that has continued and prospered for over a hundred years.<\/p>\n<p class=\"import-Normal\">Franz Boas (1858\u20121942) was a German American anthropologist who established the four-field anthropology system in the United States and founded the American Anthropological Association in 1902. He argued that the scientific method should be used in the study of human cultures and the comparative method for looking at human biology worldwide. One of Boas\u2019s significant contributions to biological anthropology was the study of skull dimensions with respect to race. After a long-term research project, he demonstrated how cranial form was highly dependent on cultural and environmental factors and that human behaviours were influenced primarily not by genes but by social learning. He wrote in one essay for the journal <em>Science<\/em>: \u201cWhile individuals differ, biological differences between races are small. There is no reason to believe that one race is by nature so much more intelligent, endowed with great willpower, or emotionally more stable than another, that the difference would materially influence its culture\u201d (Boas 1931:6). This conclusion directly contrasted with the theories of the past that relied on biological determinism. Biological anthropologists today have found evidence that corroborates Boas\u2019s explanations: societies do not exist on a hierarchy or gradation of \u201ccivilizedness\u201d but instead are shaped by the world around them, their demographic histories, and the interactions they have with other groups.<\/p>\n<figure style=\"width: 358px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image2-7.png\" alt=\"Black-and-white sketch of tree labeled \u201cEugenics.\u201d \" width=\"358\" height=\"275\" \/><figcaption class=\"wp-caption-text\">Figure 14.9: Logo of the Second International Exhibition of Eugenics, held in 1921. The text of the logo states: \"Eugenics is the self-direction of human evolution. Like a tree, eugenics draws its materials from many sources and organizes them into an harmonious entity.\" Credit: <a class=\"rId51\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Eugenics_congress_logo.png\">Eugenics congress logo<\/a> scanned from <a class=\"rId52\" href=\"https:\/\/en.wikipedia.org\/wiki\/Harry_H._Laughlin\">Harry H. Laughlin<\/a>, The Second International Exhibition of Eugenics held September 22 to October 22, 1921, is in the <a class=\"rId53\" 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 first half of the 1900s still involved some research that was essentialist and focused on proving racial determinism. Anthropologists like Francis Galton (1822\u20121911) and Earnest A. Hooton (1887\u20121954) created the field of <strong>eugenics<\/strong> as an attempt to formalize the social scientific study of \u201cfitness\u201d and \u201csuperiority\u201d among members of 19th-century Europe. As a way of \u201cdealing with\u201d criminals, diseased individuals, and \u201cuncivilized\u201d people, eugenicists recommended prohibiting parts of the population from being married or sterilizing these members of society so they could no longer procreate (Figure 14.9). They instead encouraged \u201creproduction in individual families with sound physiques, good mental endowments, and demonstrable social and economic capability\u201d (Hooton 1936). In the 1930s, Nazi Germany used this false idea of there being \u201cpure races\u201d to highly destructive effect. The need to be protected against admixture from \u201cunfit\u201d groups was their justification for their blatant racism and purging of citizens that fell under their subjective criteria.<\/p>\n<p class=\"import-Normal\">It should be noted that eugenicist ideas were popularly discussed and debated in many non-European contexts, as in the U.S., China, and South Africa, too. The Immigration Restriction Act of 1924 was passed in the United States, with the explicit aim of reducing the country\u2019s \u201cburden\u201d of people considered inferior by restricting immigration of eastern European Jews, Italians, Africans, Arabs, and Asians. In the early 1900s, Chinese scientists and politicians showed great interest in eugenic ideologies, which came to dictate decisions in law-making, family life, and the medical field. Noted American anthropologist Ruth Benedict wrote extensively on Japanese culture and society during and after World War II. Her essentialist portrayals of Japanese people were heavily cited in popular discourse at the time. In 1950s South Africa, interracial marriages and sexual relations were banned by law; antimiscegenation became one of the huge focuses of apartheid resistance activists in later years. We still see the continuation of eugenics-based logic today around the world\u2014in exclusionary immigration laws, cases of incarcerated prison inmates being forcibly sterilized, and the persistence of intelligence testing as a form of measuring people\u2019s \u201cfitness\u201d in a society.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Shortly after World War II and the Nazi Holocaust, the full extent of essentialist, eugenicist thinking became clear. Social constructions of race, and the notion that one can predict psychological or behavioural traits based on external appearance, had become unpopular both within and outside the discipline. It was up to those in the field of physical anthropology at the time to separate physical anthropology from race concepts that supported unscientific and socially damaging agendas. This does not mean that there are no physiological or behavioural differences between different members of the human species. However, going forward, a number of physical anthropologists saw human biological variation as more complicated than simple typologies could describe.<\/p>\n<h3 class=\"import-Normal\"><strong>\u201cThe New Physical Anthropology\u201d<\/strong><\/h3>\n<figure style=\"width: 219px\" class=\"wp-caption alignright\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image14-6.png\" alt=\"Black-and-white photo of a person with short hair in a white shirt and tie.\" width=\"219\" height=\"291\" \/><figcaption class=\"wp-caption-text\">Figure 14.10: Theodosius Dobzhansky, an important scientist who formulated the 20th-century \u201cmodern synthesis\u201d reconciling Charles Darwin\u2019s theory of evolution and Gregor Mendel\u2019s ideas on heredity. Credit: <a class=\"rId55\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Dobzhansky_no_Brasil_em_1943.jpg\">Dobzhansky no Brasil em 1943<\/a> by unknown photographer via <a class=\"rId56\" href=\"https:\/\/www.flickr.com\/photos\/celycarmo\/\">Flickr user Cely Carmo<\/a> is in the <a class=\"rId57\" href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">After 1950, focus steered away from the concept of \u201crace\u201d as a unit of variation and toward understanding why variation exists in <strong>population<\/strong><strong>s<\/strong> from an evolutionary perspective. This was outlined by those pioneering the \u201cnew physical anthropology,\u201d such as Sherwood Washburn, Theodosius Dobzhansky (Figure 14.10), and Julian Huxley, who borrowed this approach from contemporary population geneticists. Whether using genetic or phenotypic markers as the units of study, \u201cgroups\u201d or \u201cclusters\u201d of humans differentiated by these became defined as populations that differ in the frequency of some gene or genes. Anthropologists consider what \u201cmakes\u201d a population\u2014a group of individuals potentially capable of or actually interbreeding due to shared geographic proximity, language, ethnicity, culture, and\/or values. Put another way, a population is a local interbreeding group with reduced gene flow between themselves and other groups of humans. Members of the same population may be expected to share many genetic traits (and, as a result, many phenotypic traits that may or may not be visible outwardly).<\/p>\n<figure style=\"width: 240px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image15-6.png\" alt=\"Black-and-white photo of smiling person in a suit and tie in front of a building.\" width=\"240\" height=\"300\" \/><figcaption class=\"wp-caption-text\">Figure 14.11: Julian Huxley (1942). Credit: <a class=\"rId59\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Julian_Huxley_1-2.jpg\">Julian Huxley 1-2<\/a> by unknown photographer is in the <a class=\"rId60\" href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Thinking of humans in terms of populations was part of Julian Huxley\u2019s (1942) \u201cModern Synthesis\u201d\u2014so named because it helped to reconcile two fundamental principles about evolution that had not been made sense of together before (Figure 14.11). As discussed in Chapter 4, Gregor Mendel (1822\u20121884) was able to show that inheritance was mediated by discrete particles (or genes) and not blended in the offspring. However, it was difficult for some 19th-century scientists to accept this model of genetic inheritance at the time because much of biological variation appeared to be continuous and not particulate (take skin colour or height as examples). In the 1930s, it was demonstrated that traits could be polygenic and that multiple alleles could be responsible for any one phenotypic trait, thus producing the continuous variation in traits such as eye colour that we see today. Thus, Huxley\u2019s \u201cModern Synthesis\u201d outlines not only how human populations are capable of exchanging genes at the microevolutionary level but also how multiple alleles for one trait (polygenic exchanges) can cause gradual macroevolutionary changes.<\/p>\n<h2 class=\"import-Normal\">Human Variation in Biological Anthropology Today<\/h2>\n<h3 class=\"import-Normal\"><strong>Many Human Traits Are Nonconcordant<\/strong><\/h3>\n<p class=\"import-Normal\">In our studies of human (genetic) variation today, we understand most human traits to be nonconcordant (Figure 14.12). \u201c<strong>Nonconcordance<\/strong>\u201d is a term used to describe how biological traits vary independent of each other\u2014that is, they do not get inherited in a correlative manner with other genetically controlled traits. For example, if you knew an individual had genes that coded for tall height, you would not be able to predict if they are lighter-skinned or have red hair. This is different from earlier essentialist views of human variation: the idea that skin colour could predict one\u2019s brain function or even \u201ctemperament\u201d and tendencies toward criminal behaviour.<\/p>\n<figure style=\"width: 594px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image17-2-1.jpg\" alt=\"World map with human silhouettes scattered across the continents.\" width=\"594\" height=\"304\" \/><figcaption class=\"wp-caption-text\">Figure 14.12: Most human biological traits are non-concordant, meaning traits vary independently and each trait has its own pattern of distribution around the world. In this image, different colours and patterns represent trait varieties. For example, the colour and pattern of the head may represent hair colour (dark to light), but sharing dark hair with another person does not mean you will share other traits (e.g. ability to digest lactose or ABO blood type). Credit: Nonconcordance original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Katie Nelson is under a <a class=\"rId62\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Human Variation Is Clinal\/Continuous (Not Discrete)<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Frank B. Livingstone (1928\u20122005) wrote: \u201cThere are no races, only clines\u201d (1962: 279). A <strong>cline<\/strong> is a gradation in the frequency of an allele\/trait between populations living in different geographic regions. Human variation cannot be broken into discrete \u201craces,\u201d because most physical traits vary on a continuous or \u201cclinal\u201d basis. One obvious example of this is how human height does not only come in three values (\u201cshort,\u201d \u201cmedium,\u201d and \u201ctall\u201d) but instead varies across a spectrum of vertical heights achievable by humans all over the world. On the one hand, we can describe human height as exhibiting <strong>continuous <\/strong><strong>variation<\/strong>, forming a continuous pattern, but height does not vary according to where people live across the globe and does not exhibit a clinal pattern. On the other hand, skin colour variation between populations does show patterning that fits quite well on to how near or far they are from each other on a world map. This makes a trait like skin colour clinally distributed worldwide. When large numbers of genetic loci for large numbers of samples were sampled from human populations distributed worldwide during the 1960s and 1970s, the view that certain facets of human diversity were clinally distributed was further supported by genetic data.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">To study human traits that are clinally distributed, genetic tests must be performed to uncover the true frequencies of an allele or trait across a certain geographic space. One easily visible example of a clinal distribution seen worldwide is the patterning of human variation in skin colour. Whether in southern Asia, sub-Saharan Africa, or Australia, dark brown skin is found. Paler skin tones are found in higher-latitude populations such as those who have lived in areas like Europe, Siberia, and Alaska for millennia. Skin colour is easily observable as a phenotypic trait exhibiting continuous variation.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">A clinal distribution still derives from genetic inheritance; however, clines often correspond to some gradually changing environmental factor. Clinal patterns arise when selective pressures in one geographic area differ from those in another as well as when people procreate and pass on genes together with their most immediate neighbours. There are several mechanisms, selective and neutral, that can lead to the clinal distribution of an allele or a biological trait. <strong>Natural selection<\/strong> is the mechanism that produced a global cline of skin colour, whereby darker skin colour protects equatorial populations from high amounts of UV radiation; there is a transition of lessening pigmentation in individuals that reside further and further away from the tropics (Jablonski 2004; Jablonski and Chaplin 2000; see Figure 14.13). The ability and inability to digest lactose (milk sugar) among different world communities varies according to differential practices and histories of milk and dairy-product consumption (Gerbault Et al. 2011; Ingram Et al. 2009). Where malaria seems to be most prevalent as a disease stressor on human populations, a clinal gradient of increasing sickle cell anemia experience toward these regions has been studied extensively by genetic anthropologists (Luzzatto 2012). Sometimes, culturally defined mate selection based on some observable trait can lead to clinal variation between populations as well.<\/p>\n<figure style=\"width: 676px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image22-6.png\" alt=\"A global map shaded representing skin colors. \" width=\"676\" height=\"370\" \/><figcaption class=\"wp-caption-text\">Figure 14.13: A global map of skin colours shows that dark skin pigmentation is more common in areas that receive more UV radiation (near the equator and in high altitude areas). Light skin is more common at northern and southern latitudes. It is worth bearing in mind, though, that these do not tell the full story of how human skin pigmentation varies worldwide. Each region will contain populations that exhibit a range of skin tones. In this way, this map is not perfect as an illustration of skin-colour distribution. Credit: Mercator style projection map showing human skin colour according to Biasutti 1940.png by Dark Tichondrias at English Wikipedia, modified (cropped) by Tuvalkin, is under a CC BY-SA 4.0 License.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Two neutral microevolutionary processes that may produce a cline in a human allele or trait are <strong>gene flow<\/strong> and <strong>genetic drift <\/strong>(see Chapter 5). The ways in which neutral processes can produce clinal distributions is seen clearly when looking at clinal maps for different blood groups in the human ABO blood group system (Figure 14.14). For instance, scientists have identified an East-to-West cline in the distribution of the blood type <em>B<\/em> allele across Eurasia. The frequency of <em>B<\/em> allele carriers decreases gradually westward when we compare the blood groups of East and Southeast Asian populations with those in Europe. This shows how populations residing nearer to one another are more likely to interbreed and share genetic material (i.e., undergo gene flow). We also see 90%\u2012100% of native South American individuals, as well as between 70%\u201290% of Aboriginal Australian groups, carrying the <em>O<\/em> allele (Mourant, \u200b\u200bKope\u0107, and Domaniewska-Sobczak 1976). These high frequencies are likely due to random genetic drift and founder effects, in which population sizes were severely reduced by the earliest <em>O<\/em> allele-carrying individuals migrating into those areas. Over time, the <em>O<\/em> blood type has remained predominant.<\/p>\n<p class=\"import-Normal\"><img class=\"aligncenter\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image8.gif\" alt=\"World map showing varying frequencies of blood type A.\" width=\"516\" height=\"284\" \/><\/p>\n<p class=\"import-Normal\"><img class=\"aligncenter\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5.gif\" alt=\"World map shows the highest frequencies of blood type B in parts of Asia.\" width=\"518\" height=\"283\" \/><\/p>\n<figure style=\"width: 516px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image13.gif\" alt=\"World map shows the highest frequencies of blood type O.\" width=\"516\" height=\"284\" \/><figcaption class=\"wp-caption-text\">Figure 14.14a\u2013c: a. Global distribution of blood type A. b. Global distribution of blood type B. c. Global distribution of blood type O. <a href=\"https:\/\/docs.google.com\/document\/d\/1VUDKMBJYS_jNONjLxT04jQN0_z9Ua50BRN6auGSHUuU\/edit\" target=\"_blank\" rel=\"noopener\">A text description of this image is available<\/a>. Credit: a. <a class=\"rId69\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Map_of_blood_group_a.gif\">Map of blood group a<\/a> by <a class=\"rId70\" href=\"https:\/\/en.wikipedia.org\/wiki\/User:Muntuwandi\">Muntuwandi<\/a> at <a class=\"rId71\" href=\"https:\/\/en.wikipedia.org\/\">en.wikipedia<\/a> is under a <a class=\"rId72\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>. b. <a class=\"rId73\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Map_of_blood_group_b.gif\">Map of blood group b<\/a> by <a class=\"rId74\" href=\"https:\/\/en.wikipedia.org\/wiki\/User:Muntuwandi\">Muntuwandi<\/a> at <a class=\"rId75\" href=\"https:\/\/en.wikipedia.org\/\">en.wikipedia<\/a> is under a <a class=\"rId76\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>. c. <a class=\"rId77\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Map_of_blood_group_o.gif\">Map of blood group o<\/a> is based on diagrams from <a class=\"rId78\" href=\"https:\/\/anthro.palomar.edu\/vary\/vary_3.htm\">https:\/\/anthro.palomar.edu\/vary\/vary_3.htm<\/a>, reproduced from A. E. Mourant Et al. (1976), and is under a <a class=\"rId79\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Genetic Variation Is Greater Within Group than Between Groups<\/strong><\/h3>\n<figure style=\"width: 295px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image27-2.jpg\" alt=\"Two medium-sized circles labeled Asian and European largely overlap. A larger circle labeled African surrounds both. \" width=\"295\" height=\"274\" \/><figcaption class=\"wp-caption-text\">Figure 14.15: Circles represent human genetic variation. Most variants are shared among individuals on all continents. There are more variants in Africa, some of which are not found in Europe or Asia. Credit: Human Genetic Variation original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Katie Nelson is under a <a class=\"rId81\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p>One problem with race-based classifications is they relied on an erroneous idea that individuals with particular characteristics would share more similar genes with each other within a particular \u201crace\u201d and share less with individuals of other \u201craces\u201d possessing different traits and genetic makeups. However, since around 50 years ago, scientific studies have shown that the majority of human genetic differences worldwide exist <em>within<\/em> groups (or \u201craces\u201d) individually rather than <em>between<\/em> groups. Indeed, most genetic variation we see occurs in Africa, and many variants are shared among individuals on all continents (Figure 14.15).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In 2002, a landmark article by Noah Rosenberg and colleagues explored worldwide human genetic variation using an even-greater genetic data set. They used 377 highly variable markers in the human genome and sampled from 1,056 individuals representative of 52 populations. The markers chosen for study were not ones that code for any expressed genes. Because these regions of the human genome were made of unexpressed genes, we may understand these markers as neutrally derived (as opposed to selectively derived) because they do not code for functional advantages or disadvantages. These neutral genetic markers likely reflect an intricate combination of regional founder effects and population histories. Analyses of these neutral markers allowed scientists to identify that 93%\u201295% of global genetic differences, referred to as <strong>variance<\/strong>, can be accounted for by within-population differences, while only a small proportion of genetic variance (3%\u20125%) can be attributed to differences among major groups (Rosenberg Et al. 2002). This research supports the theory that distinct biological races do not exist, even though misguided concepts of race may still have real social and political consequences.<\/p>\n<h3 class=\"import-Normal\"><strong>Biological Data Fit Isolation-By-Distance and Out-of-Africa Models<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">One further note is that the world\u2019s population may be genetically divided into \u201cgroups,\u201d \u201csubsets,\u201d \u201cclumps,\u201d or \u201cclusters\u201d that reflect some degree of genetic similarity. These identifiable clusters reflect genetic or geographic distances\u2014either with gene flow facilitated by proximity between populations or impeded by obstacles like oceans or environmentally challenging habitats (Rosenberg Et al. 2005). Sometimes, inferred clusters using multiple genetic loci are interpreted by nongeneticists literally as \u201cancestral populations.\u201d However, it would be wrong to assume from these genetic results that highly differentiated and \u201cpure\u201d ancestral groups ever existed. These groupings reflect differences that have arisen over time due to clinal patterning, genetic drift, and\/or restricted or unrestricted gene flow (Weiss and Long 2009). The clusters identified by scientists are arbitrary and the parameters used to split up the global population into groups is subjective and dependent on the particular questions or distinctions being brought into focus (Relethford 2009).<\/p>\n<figure style=\"width: 341px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image6-7.png\" alt=\"Map of the continent of Africa with the lower two-thirds shaded.\" width=\"341\" height=\"341\" \/><figcaption class=\"wp-caption-text\">Figure 14.16: Sub-Saharan Africa (shaded dark\/green). Credit: <a class=\"rId83\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Sub-Saharan-Africa.png\">Sub-Saharan Africa<\/a> by <a class=\"rId84\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Ezeu\">Ezeu<\/a> has been designated to the <a class=\"rId85\" 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\">Additionally, research on worldwide genetic variation has shown that human variation decreases with increasing distance from sub-Saharan Africa, where there is evidence for this vast region being the geographical origin of anatomically modern humans (Liu Et al. 2006; Prugnolle, Manica, and Balloux 2005; see Figures 14.16 and 14.17). Genetic differentiation decreases in human groups the further you sample data from relative to sub-Saharan Africa because of serial founder effects (Relethford 2004). Over the course of human colonization of the rest of the world outside Africa, populations broke away in expanding waves across continents into western Asia, then Europe and eastern Asia, followed by Oceania and the Americas. As a result, founder events occurred whereby genetic variation was lost, as the colonization of each new geographical region involved a smaller number of individuals moving from the original larger population to establish a new one (Relethford 2004). The most genetic variation is found across populations residing in different parts of sub-Saharan Africa, while other current populations in places like northern Europe and the southern tip of South America exhibit some of the least genetic differentiation relative to all global populations (Campbell and Tishkoff 2008).<\/p>\n<figure style=\"width: 469px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image3-6.png\" alt=\"Two scatterplots.\" width=\"469\" height=\"516\" \/><figcaption class=\"wp-caption-text\">Figure 14.17: Comparison of the genetic distance and geographical distance between populations. In the top graph, the pattern reveals that genetic variation conforms to an Out-of-Africa model, as those populations further away from Addis Ababa in Ethiopia share a smaller number of alleles; in the bottom graph, we see the populations follow an isolation-by-distance model, as pairs of populations further apart geographically seem to have greater genetic distance (Kanitz Et al. 2018). Credit: <a class=\"rId87\" href=\"https:\/\/journals.plos.org\/plosone\/article?id=10.1371\/journal.pone.0192460\">Complex genetic patterns (figure 1)<\/a> by Kanitz Et al. (2018) is under a <a class=\"rId88\" href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/\">CC BY 4.0 License<\/a>.<\/figcaption><\/figure>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<figure style=\"width: 377px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image1-9.png\" alt=\"Two large circles connected by a small area.\" width=\"377\" height=\"309\" \/><figcaption class=\"wp-caption-text\">Figure 14.18: The founder effect is a change in a small population\u2019s gene pool due to a limited number of individuals breaking away from a parent population. Credit: <a class=\"rId90\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Bottleneck_effect.jpg\">Bottleneck effect<\/a> by <a class=\"rId91\" href=\"https:\/\/wikieducator.org\/User:Tsaneda\">Tsaneda<\/a> is under a <a class=\"rId92\" href=\"https:\/\/creativecommons.org\/licenses\/by\/3.0\/legalcode\">CC BY 3.0 License<\/a>.<\/figcaption><\/figure>\n<p>Besides fitting nicely into the <strong>Out-of-Africa model<\/strong>, worldwide human genetic variation conforms to an <strong>isolation-by-distance model<\/strong>, which predicts that genetic similarity between groups will decrease exponentially as the geographic distance between them increases (Kanitz Et al. 2018). This is because of the greater and greater restrictions to gene flow presented by geographic distance, as well as cultural and linguistic differences that occur as a result of certain degrees of isolation. Since genetic data conform to isolation-by-distance and Out-of-Africa models, these findings support the abolishment of \u201crace\u201d groupings. This research demonstrates that human variation is continuous and cannot be differentiated into geographically discrete categories. There are no \u201cinherent\u201d or \u201cinnate\u201d differences between human groups; instead, variation derives from some degree of natural selection, as well as neutral processes like <strong>population bottle-necking<\/strong> (Figure 14.18), random <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1474\"><strong>mutations<\/strong><\/a> in the DNA, genetic drift, and gene flow through between-mate interbreeding.<\/p>\n<h3 class=\"import-Normal\"><strong>Humans Have Higher Homogeneity Compared to Many Other Species<\/strong><\/h3>\n<p class=\"import-Normal\">An important fact to bear in mind is that humans are 99.9% identical to one another. This means that the apportionments of human variation discussed above only concern that tiny 0.1% of difference that exists between all humans globally. Compared to other mammalian species, including the other great apes, human variation is remarkably lower. This may be surprising given that the worldwide human population has already exceeded seven billion, and, at least on the surface level, we appear to be quite phenotypically diverse. Molecular approaches to human and primate genetics tells us that external differences are merely superficial. For a proper appreciation of human variation, we have to look at our closest relatives in the primate order and mammalian class. Compared to chimpanzees, bonobos, gorillas and other primates, humans have remarkably low average genome-wide <strong>heterogeneity<\/strong> (Osada 2005).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">When we look at chimpanzee genetic variation, it is fascinating that western, central, eastern, and Cameroonian chimpanzee groups have substantially more genetic variation between them than large global samples of human DNA (Bowden Et al. 2012; Figure 14.19). This is surprising given that all of these chimpanzee groups live relatively near one another in Africa, while measurements of human genetic variation have been conducted using samples from entirely different continents. First, geneticists suppose that this could reflect differential experiences of the founder effect between humans and chimpanzees. As it has been argued that all non-African human populations descended from a small number of anatomically modern humans who left Africa, it would be expected that all groups descended from that smaller ancestral group would be similar genetically. Second, our species is really young, given that we have only existed on the planet for around 150,000 to 300,000 years. This gave humans little time for random genetic mutations to occur as genes get passed down through genetic interbreeding and meiosis. Chimpanzees, however, have inhabited different <strong>ecological niches<\/strong>, and less interbreeding has occurred between the four chimpanzee groups over the past six to eight million years compared to the amount of gene flow that occurred between worldwide human populations (Bowden Et al. 2012).<\/p>\n<figure style=\"width: 648px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image16-7.png\" alt=\"Map of Africa showing the ranges of chimpanzees from west to east.\" width=\"648\" height=\"339\" \/><figcaption class=\"wp-caption-text\">Figure 14.19: Distribution of the genus Pan, including bonobos and the four subspecies of chimpanzee, across western and central Africa (Clee Et al. 2015). <a href=\"https:\/\/docs.google.com\/document\/d\/1VUDKMBJYS_jNONjLxT04jQN0_z9Ua50BRN6auGSHUuU\/edit\" target=\"_blank\" rel=\"noopener\">A text description of this image is available<\/a>. Credit: <a class=\"rId94\" href=\"https:\/\/bmcecolevol.biomedcentral.com\/articles\/10.1186\/s12862-014-0275-z\">Chimpanzee subspecies ranges (Figure 1)<\/a> by Clee Et al. 2015 is under a <a class=\"rId95\" 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\">Recent advances have now enabled the attainment of genetic samples from the larger family of great apes and the evaluation of genetic variation among bonobos, orangutans, and gorillas alongside that of chimpanzees and humans (Prado-Martinez Et al. 2013). Collecting such data and analyzing primate genetic variation has been important not only to elucidate how different ecological, demographic, and climatic factors have shaped our evolution but also to inform upon conservation efforts and medical research. Genes that may code for genetic susceptibilities to tropical diseases that affect multiple primates can be studied through genome-wide methods. Species differences in the genomes associated with speech, behaviour, and cognition could tell us more about how human individuals may be affected by genetically derived neurological or speech-related disorders and conditions (Prado-Martinez Et al. 2013; Staes Et al. 2017). In 2018, a great ape genomic study also reported genetic differences between chimpanzees and humans related to brain cell divisions (Kronenberg Et al. 2018). From these results, it may be inferred that cognitive or behavioural variation between humans and the great apes might relate to an increased number of cortical neurons being formed during human brain development (Kronenberg Et al. 2018). Comparative studies of human and nonhuman great ape genetic variation highlight the complex interactions of population histories, environmental changes, and natural selection between and within species. When viewed in the context of overall great ape variation, we may reconsider how variable the human species is relatively and how unjustified previous \u201crace\u201d concepts really were.<\/p>\n<h3 class=\"import-Normal\"><strong>Phenotypic Traits That Reflect Neutral Evolution<\/strong><\/h3>\n<p class=\"import-Normal\">Depending on the trait being observed, different patterns of phenotypic variation may be found within and among groups worldwide. In this subsection, some phenotypic traits that reflect the aforementioned patterns of genetic variation will be discussed.<\/p>\n<figure style=\"width: 301px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image18-4.jpg\" alt=\"Illustration of one anthropologist studying teeth, and another looking into a microscope.\" width=\"301\" height=\"301\" \/><figcaption class=\"wp-caption-text\">Figure 14.20: Contemporary anthropologists who use many types of skeletal markers have demonstrated that a majority of cranial variation occurs within populations rather than between populations and that there is a decrease in variation with distance from Africa. Credit: <a class=\"rId97\" href=\"https:\/\/img1.wsimg.com\/isteam\/ip\/0f6c1c17-41ea-4caf-b839-73c676d69f01\/DentalHeartNecklace.jpg\/:\/rs=w:1280,h:1280\">Dental Anthropologist Heart Necklace<\/a> by <a class=\"rId98\" href=\"https:\/\/anthroillustrated.com\/\">Anthro Illustrated<\/a> is under a <a class=\"rId99\" 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\">Looking beyond genetic variation briefly, recent studies have revisited biological anthropology\u2019s earlier themes of externally observable traits, such as skull shape. In the last 20 or so years, anthropologists have evaluated the level to which human cranial shape variation reflects the results from genetic markers, such as those used previously to fit against Out-of-Africa models (Relethford 2004) or those used in the apportionment of human variation between and within groups (Lewontin 1972; Rosenberg Et al. 2002). Using larger sample sizes of cranial data collected from thousands of skulls worldwide and a long list of cranial measurements, studies demonstrate a similar decrease in variation with distance from Africa and show that a majority of cranial variation occurs within populations rather than between populations (Betti Et al. 2009; Betti Et al. 2010; Manica Et al. 2007; Relethford 2001; von Cramon-Taubadel and Lycett 2008; see Figure 14.20). The greatest cranial variation is found among skulls of sub-Saharan African origin, while the least variation is found among populations inhabiting places like Tierra del Fuego at the southern tip of Argentina and Chile. While ancient and historical thinkers previously thought \u201crace\u201d categories could reasonably be determined based on skull dimensions, modern-day analyses using more informative sets of cranial traits simply show that migrations out of Africa and the relative distances between populations can explain a majority of worldwide cranial variation (Betti Et al. 2009).<\/p>\n<figure style=\"width: 250px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image21-6.png\" alt=\"Sketch of bone with bony loops at the top and coil at the bottom.\" width=\"250\" height=\"208\" \/><figcaption class=\"wp-caption-text\">Figure 14.21: Diagram of the bony labyrinth in the inner ear. Credit: <a class=\"rId101\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Bony_labyrinth.png\">Bony labyrinth<\/a> by <a class=\"rId102\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Selket\">Selket<\/a> has been designated to the <a class=\"rId103\" href=\"https:\/\/creativecommons.org\/share-your-work\/public-domain\/cc0\/\">public domain (CC0)<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">This same patterning in phenotypic variation has even been found in studies examining shape variation of the pelvis (Betti Et al. 2013; Betti Et al. 2014), the teeth (Rathmann Et al. 2017), and the human <strong>bony labyrinth<\/strong> of the ear (Ponce de Le\u00f3n Et al. 2018;Figure 14.21). The skeletal morphology of these bones still varies worldwide, but a greater proportion of that variation can still be attributed to the ways in which human populations migrated across the world and exchanged genes with those closer to them rather than those further away. Human skeletal variation in these parts of the body is continuous and nondiscrete. Given the important functions of the cranium and these other skeletal parts, we may infer that the genes that underpin their development have been relatively conserved by neutral evolutionary processes such as genetic drift and gene flow. It is also important to note that while some traits such as height, weight, cranial dimensions, and body composition are determined, in part, by genes, the underlying developmental processes behind these traits are underpinned by complex polygenic mechanisms that have led to the continuous spectrum of variation in such variables among modern-day human populations.<\/p>\n<h3 class=\"import-Normal\"><strong>Phenotypic Traits That Reflect Natural Selection<\/strong><\/h3>\n<p class=\"import-Normal\">Even though 99.9% of our DNA is the same across all humans worldwide, and many traits reflect neutral processes, there are parts of that remaining 0.1% of the human genome that code for individual and regional differences. Similarly to craniometric analyses that have been conducted in recent decades, human variation in skin colour has also been reassessed using new methods and in light of greater knowledge of biological evolution.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">New technologies allow scientists to use colour photometry to sample and quantify the visible wavelength of skin colour, in a way 19th- and 20th-century readers could not. In one report, it was found that 87.9% of global skin colour variation can be attributed to genetic differences <em>between<\/em> groups, 3.2% to those among local populations within regions, and 8.9% <em>within<\/em> local populations (Relethford 2002). This apportionment differs significantly and is the reverse situation found in the distribution of genetic differences we see when we examine genetic markers such as blood type\u2013related alleles. However, this pattern of human skin colour worldwide is not surprising, given that we now understand that past selection has occurred for darker skin near the equator and lighter skin at higher latitudes (Jablonski 2004; Jablonski and Chaplin 2000). While most genetic variation reflects neutral variation due to population migrations, geographic isolation, and restricted gene flow dynamics, some human genetic\/phenotypic variation is best explained as local adaptation to environmental conditions (i.e., selection). Given that skin colour variation is atypical compared to other genetic markers and biological traits, this, in fact, goes against earlier \u201crace\u201d typologies. This is because recent studies ironically show how so much of genetic variation relates to neutral processes, while skin colour does not. It follows that skin colour <em>cannot<\/em> be viewed as useful in making inferences about other human traits.<\/p>\n<figure style=\"width: 580px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image23-3.jpg\" alt=\"Person standing at podium in front of a screen with arms partly raised.\" width=\"580\" height=\"384\" \/><figcaption class=\"wp-caption-text\">Figure 14.22: Genomicists and biological anthropologists have dedicated efforts to improving quantitative methods of measuring hair and skin variation over the last twenty years. Dr. Nina Jablonski is one such biological anthropologist specializing in the evolution and variation of human skin pigmentation. Credit: <a class=\"rId105\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Nina_Jablonski_2016_The_Skin_of_Homo_sapiens_01_%28cropped%29.jpg\">Nina Jablonski 2016 The Skin of Homo sapiens 01 (cropped)<\/a> by <a class=\"rId106\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Ptolusque\">Ptolusque<\/a> is under a <a class=\"rId107\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\">CC BY-SA 4.0 license<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">It is also true that some populations have not been studied extensively in skin pigmentation genetics (e.g., African, Austronesian, Melanesian, Southeast Asian, Indigenous American, and Pacific Islander populations, according to Lasisi and Shriver 2018). Earlier dispersals of these populations, and their local genetic varition, will have contributed to worldwide genetic variation, inclusive of skin pigmentation variation. Gene loci we did not previously appreciate as being linked to pigmentation are now being recognized thanks to better tools, more diverse genetic samples, and more accessible datasets (Quillen Et al. 2018). Biological anthropologists look forward to further discoveries elucidating the different selective pressures and population dynamics that influence skin pigmentation evolution.<\/p>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Social Implications<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">To finish this chapter, we will consider the social, economic, political, and biological implications of poor understandings of race and the deliberate perpetuation of social and medical racism.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The Black Lives Matter movement (BLM) of 2013 began with the work of racial justice activists and community organizers Alicia Garza, Opal Tometi, and Patrissa Cullors. First incited by the murder of Trayvon Martin, a 17-year-old African American, and the acquittal of the man who shot him, BLM went on to protest against the deaths of numerous Black individuals, most of whom were killed by police officers (for example, Ahmaud Arbery was killed in February of 2020 by two white non-police officers). Some key characteristics of BLM from the start were its decentralized grassroots structure, the role of university students and social media in spreading awareness of the movement, and its embrace of other movements (e.g., climate justice, ending police brutality, feminist campaigns, queer activism, immigration reform, etc.). When George Floyd was murdered by a white police officer on May 25, 2020, the BLM gained new momentum, across 2,000-plus cities in the United States, and among many protesting against historic racism and police brutality in other contexts around the globe. Many in the biological anthropology community have responded to these events with a great dedication to working against systemic racism in society and institutions (American Association of Biological Anthropologists 2020).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">BLM continues to be an important movement, as is evidenced in the degree of community organizing, mutual aid efforts, calls for political reform, progress toward curriculum reform and equality, inclusion and diversity (EDI) work in businesses and universities, the removal of monuments honoring historical figures associated with slavery and racism, and many other important actions. Garza (2016) writes: \u201cThe reality is that race in the United States operates on a spectrum from black to white \u2026 the closer you are to white on that spectrum, the better off you are.\u201d Tometi (2016) has stated: \u201cWe need [a human rights movement that challenges systemic racism] because the global reality is that Black people are subject to all sorts of disparities in most of our challenging issues of our day. I think about climate change, and how six of the ten worst impacted nations by climate change are actually on the continent of Africa.\u201d In the words of Cullors (2016), \u201cBlack Lives Matter is our call to action. It is a tool to reimagine a world where Black people are free to exist, free to live. It is a tool for our allies to show up differently for us.\u201d We gather from their words the importance of learning from the egregious role that anthropologists have played in the past, recognizing the legacies of \u201cscientific\u201d justifications for eugenics and racism in our society today, and proactively working toward environmental and social equity.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Another major industry that engages in the quantification and interpretation of human variation is medical and clinical work (National Research Council [U.S.] Committee on Human Genome Diversity 1997). Large-scale genomic studies sampling from human populations distributed worldwide have produced detailed knowledge on variation in disease resistance or susceptibility between and within populations. Let\u2019s think about drug companies who develop medicines for Black patients particularly. The predispositions to particular diseases are higher among people of African descent than some pharmaceutical businesses have taken into account. Through targeted sampling of various world groups, clinical geneticists may also identify genetic risk factors of certain common disorders such as chronic heart disease, asthma, diabetes, autoimmune diseases, and behavioural disorders. Having an understanding of population-specific biology is crucial in the development of therapies, medicines, and vaccinations, as not all treatments may be suitable for every human, depending on their genotype. During diagnosis and treatment, it is important to have an evolutionary perspective on gene-environment relationships in patients. Typological concepts of \u201crace\u201d are not useful, given that most racial groups (whether self-identified or not) popularly recognized lack homogeneity and are, in fact, variable. <strong>Cystic fibrosis<\/strong>, for instance, occurs in all world populations but can often be underdiagnosed in populations with African ancestry because it is thought of as a \u201cwhite\u201d disease (Yudell Et al. 2016).<\/p>\n<p class=\"import-Normal\">Sociologists, law scholars, and professors of race studies have written extensively on how genetic\/technological\/medical revolutions impact people of colour. In her book, <em>Fatal Invention: How Science, Politics, and Big Business Re-create Race in the Twenty-First Century <\/em>(2013), Professor Dorothy E. Roberts writes about how technological advances have been used in resuscitating race as a biological category for dividing humans in essentialist ways (Figure 14.23). She notes how members of law enforcement have engaged in racial profiling, sometimes with the use of machine-learning and facial-recognition technologies. Ancestry-testing services also purport to tell us \u201cwhat\u201d we are and to insist that this information is \u201cwritten\u201d in our genes. Such advertising campaigns obscure the nuances of genetic variation with the primary motive of tapping into people\u2019s desire to \u201cknow themselves\u201d and driving up profits for their businesses. Commercial genetic testing reinforces the idea that genes map neatly onto race, all while generating massive stores of data in DNA databases. In Roberts\u2019s view, the myth of the biological concept of race being perpetuated in these ways undermines a just society and reproduces racial inequalities.<\/p>\n<figure style=\"width: 593px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image12-6.png\" alt=\"Book cover with two DNA strands and next to the smiling portrait of the author.\" width=\"593\" height=\"305\" \/><figcaption class=\"wp-caption-text\">Figure 14.23: Professor Dorothy E. Roberts is a sociologist, legal scholar, and expert on the relationships among technology, medicine, bioethics, policymaking, race, and racism. Credit: <a class=\"rId109\" href=\"https:\/\/kpfa.org\/episode\/talkies-august-23-2016\/\">Dorothy Roberts author of Fatal Invention<\/a> by <a class=\"rId110\" href=\"https:\/\/kpfa.org\/\">https:\/\/kpfa.org\/<\/a> is copyrighted and used with permission.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The COVID-19 pandemic has had a significant impact on the world\u2019s population, particularly people living in the economic Global South and many Black, Indigenous and communities of colour residing in the Global North. We have witnessed disproportionately high numbers of COVID-related deaths and infection cases among marginalized groups. Many immigrants and ethnic minorities in various societies have also experienced scapegoating and blame directed at them for being the source of COVID-19 spread.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">To inform us on how to interpret this current worldwide pandemic, historians and anthropologists are looking back at the lessons learned from past instances of racist medicine (discriminatory practices based on broader social discrimination) and medical racism (application of discriminatory practices justified on medical grounds). Historically, who could become doctors and medical professionals was often racialized, gendered, and class specific. This made it difficult for many to overcome prejudices against women, Black people, Indigenous individuals, or other people of colour from becoming doctors and clinical researchers in places such as South Africa and the United States. This, in turn, affects the sorts of information we know about health levels and health outcomes among these very groups. In the past decade, long-overdue attention is finally being paid to how race affects biological outcomes. For instance, researchers have focused on the negative legacies of racial discrimination and racism-induced stress on hormone (im)balances, mental health disorders, cardiovascular disease prevalence, and other health outcomes (Kuzawa and Sweet 2009; Shonkoff, Slopen, and WIlliams 2021; Williams 2018). The technology and standards of protocol in medical testing have been scrutinized (for more on how pulse oximeters were not designed with nonwhite patients in mind, for example, see Sjoding Et al. 2020). Scholars of race and medicine have also written on how illness and disease spread have often been used to perpetuate societal prejudices. This manifests as xenophobic tendencies at a societal level, such as the blaming of \u201coutgroups\u201d and increased \u201cin-group\u201d protectiveness. Overreliance on the idea that people are \u201cinherently\u201d disease carriers due to genetic or biological reasons leads to improper accounting for socioeconomic or infrastructural issues that lead to differential disease prevalence amongst minority communities. (For more on race and COVID, see Tsai 2021 as well as this textbook\u2019s Chapter 16: Contemporary Topics: Human Biology and Health.)<\/p>\n<p>Lastly, consider the changing field of forensic anthropology. In the past, forensic anthropologists ascribed <strong>ancestry<\/strong> or racial categories to sets of skeletons, reliant on the belief that different human groups will exhibit biologically \u201cdiscrete\u201d assortments so as to divide along culturally constructed categories (Sauer 1992). Now, a number of forensic anthropologists have argued that we should abandon these methods, both because it is unscientific and because it further validates and perpetuates this idea that race is biologically meaningful. As scientists, whether we affirm biological race as real has huge influence on the beliefs of members of the public, the judicolegal system, and law enforcement. Not all forensic experts agree with abandoning ancestry estimation. Some prefer to refocus on the neutral or selective <em>causes<\/em> of human biological variation, and assess how <em>probabilistic<\/em> it may be to assign bones of certain dimensions to one of several identified racial categories. These debates continue today as this textbook chapter is being written. More details on population affinity may be found in Chapter 17: Forensic Anthropology.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">It is important to remember that while it is possible to look for clues about one\u2019s ancestry or geographic origin based on skull morphology, again, the amount of distinctiveness in any given sample makes it impossible to distinguish whether a cranium belongs to one group (Relethford 2009). Individuals can vary in their skeletal dimensions by continental origin, country origin, regional origin, sex, age, environmental factors, and the time period in which they lived, making it difficult to assign individuals to particular categories in a completely meaningful way (Ousley, Jantz, and Freid 2009). When forensic reports and scientific journal articles give an estimation of ancestry, it is crucial to keep in mind that responsible assignments of ancestry will be done through robust statistical testing and stated as a probability estimate. Today, we also live in a more globalized world where a skeletal individual may have been born originally to parents of two separate traditional racial categories. In contexts of great heterogeneity within populations, this definitely adds difficulty to the work of forensic scientists and anthropologists preparing results for the courtroom (genetic testing may be comparatively more helpful in such situations).<\/p>\n<\/div>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Dig Deeper: Measuring F<sub>ST<\/sub><\/h2>\n<p class=\"import-Normal\">Richard Lewontin (1929\u2012) is a biologist and evolutionary geneticist who authored an article evaluating where the total genetic variation in humans lies. Titled \u201cThe Apportionment of Human Diversity\u201d (Lewontin 1972), the article addressed the following question: On average, how genetically similar are two randomly chosen people from the same group when compared to two randomly chosen people from different groups?<\/p>\n<p class=\"import-Normal\">Lewontin studied this problem by using genetic data. He obtained data for a large number of different human populations worldwide using 17 genetic markers (including alleles that code for various important enzymes and proteins, such as blood-group proteins). The statistical analysis he ran used a measure of human genetic differences in and among populations known as the fixation index (F<sub>ST<\/sub>).<\/p>\n<p class=\"import-Normal\">Technically, F<sub>ST<\/sub> can be defined as the proportion of total genetic variance within a <em>subpopulation<\/em> relative to the total genetic variance from an <em>entire population<\/em>. Therefore, F<sub>ST<\/sub> values range from 0 to 1 (or, sometimes you will see this stated as a percentage between 0% and 100%). The closer the F<sub>ST<\/sub> value of a population (e.g., the world\u2019s population) approaches 1, the higher the degree of genetic differentiation among subpopulations relative to the overall population (see Figure 14.24 for a detailed illustration).<\/p>\n<figure style=\"width: 561px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-5.jpg\" alt=\"Three cases each illustrate two populations with a mix of two types of alleles.\" width=\"561\" height=\"473\" \/><figcaption class=\"wp-caption-text\">Figure 14.24: This diagram shows a range of different case studies with which we may understand how FST is calculated in different populations. In Case 1, the gene pools of Populations 1 and 2 are 100% different from each other but possess 0% variation within themselves, so FST has a value of 1. When there is no genetic variation at all between two populations and 100% variation within them, as in Case 2, we see that FST is calculated as 0. When we look at Case 3, where variation between and within are some values between 0% and 100%, we will get a decimal figure for FST dependent upon how much variation there is between and within populations. It is through such comparisons of population genetic data that we may quantify the relative similarities or differences between and within populations, and we may thus speak to the nonexistence of \u201cracial groups\u201d that divide up our species into broad continental or racial categories. Credit: F<sub>ST<\/sub> original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Katie Nelson is under a <a class=\"rId112\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p>&nbsp;<\/p>\n<p>In his article, Lewontin (1972) identified that most of human genetic differences (85.4%) were found within local subpopulations (e.g., the Germans or Easter Islanders), whereas 8.3% were found between populations within continental human groups, and 6.3% were attributable to traditional \u201crace\u201d groups (e.g., \u201cCaucasian\u201d or \u201cAmerind\u201d). These findings have been important for scientifically rejecting the existence of biological races (Long and Kittles 2003).<\/p>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<h2 class=\"import-Normal\">Talking About Human Biological Variation Going Forward<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">To conclude, utilizing the term <em>races<\/em> to describe human biological variation is not accurate or productive. Using a select few hundred genetic loci, or perhaps a number of phenotypic traits, it may be possible to assign individuals to a geographic ancestry, but what constitutes a bounded genetic or geographical grouping is both arbitrary and potentially harmful owing to ethical and historical reasons. The discipline of biological anthropology has moved past typological frameworks that shoehorn continuously variable human populations into discrete and socially constructed subsets. Improvements in the number of markers, the genetic technologies used to study variation, and the number of worldwide populations sampled have led to more nuanced understandings of human variation. It is of utmost importance that scientists make the following points clear to the public:<\/p>\n<ul>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Today, we refer to different local human groups as \u201cpopulations.\u201d What constitutes a population should be carefully defined in scientific reports based on some geographical, linguistic, or cultural criteria and some degree of relativity to other closely or distantly related human groups.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Humans have significantly less genetic variation than other primates and mammals, and all human beings on Earth share 99.9% of their overall DNA. Some of the remaining 0.1% of human variation varies on a clinal or continuous basis, such as can be seen when looking at ABO blood-type <strong>polymorphisms<\/strong> worldwide.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Many biological characteristics in humans are actually determined nonconcordantly and\/or polygenically. Therefore, superiority or inferiority in human behaviour or body form cannot justifiably be linked to fixed and innate differences between groups.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Genetic distances are correlated with geographic distances among the global human population. This is especially apparent when we consider that genetic variation is highest in sub-Saharan Africa, and average genetic heterogeneity decreases in populations further away from the African continent in accordance with the migratory history of anatomically modern <em>Homo sapiens<\/em>.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">The effects of gene flow, genetic drift, and population bottlenecking are reflected in some phenotypic traits, such as cranial shape.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">We recognize other traits, like skin colour and lactase persistence, to be the products of many millennia of natural selective pressures influencing human biology from the external environment.<\/li>\n<\/ul>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Taken together, genetic analyses of human variation do not support 20th-century (or even-earlier) concepts of race. In discussions about human variation, these genomic results help clarify how biological variation is distributed across the human population today. Taking care to think about and debate the nature of human variation is important, because although the effects and events that produced genetic differences among groups occurred in the ancient past, sociocultural concepts about race and ethnicity continue to have real social, economic, and political consequences.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Beyond talking about variation in the university setting, it is important that teachers, researchers, and students of anthropology recognize and assume the responsibility of influencing public perspectives of human variation. Race-based classification systems were developed during the colonial era, the transatlantic trafficking of kidnapped Africans and the so-called \u201cScientific Revolution\u201d by the first \u201canthropologists\u201d and scholars of humankind\u2019s variation. Unfortunately, some of their early ideas have persisted and evolved into present-day lived realities. Some of today\u2019s politicians and socioeconomic bodies have racially charged agendas that promote racism or certain kinds of economic or racial inequalities. As anthropologists, we must acknowledge that while human \u201craces\u201d are not a biological reality, their status as a (misguided) social construction does have real consequences for many people (Antrosio 2011).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In other words, while \u201crace\u201d is a sociocultural invention, the treatment different individuals receive due to their perceived \u201crace\u201d can have significant financial, emotional, sociopolitical, and physiological costs. However\u2014and importantly assuming a \u201ccolour-blind\u201d position when it comes to the topics of \u201crace\u201d and ethnicity (especially in political discussions) is actually counterproductive, because the negative social consequences of modern \u201crace\u201d ideas could be ignored, making it harder to examine and address instances of discrimination properly (Wise 2010). Rather than shy away from these topics, we can use our scientific findings to establish socially relevant and biologically accurate ideas concerning human diversity. Today, research into genetic and phenotypic differentiation among and within various human populations continues to expand in its scope, its technological capabilities, its sample sizes, and its ethical concerns. It is thanks to such scientific work done in the past few decades that we now have a deeper understanding not only of how humans vary but also of how we are biologically a rather homogenous, intermixing world population.<\/p>\n<p>&nbsp;<\/p>\n<div class=\"textbox shaded\">\n<h2>Summary<\/h2>\n<p>Historically, most concepts of race were shaped by religious and early \u2018scientific\u2019 attempts to classify people, many of which justified inequality and racism. Today, biological anthropology emphasizes that human variation is continuous (clinal), polygenic, and shaped by both evolutionary pressures and neutral processes. Misunderstandings of race, however, continue to have serious social and medical consequences; evident in systemic racism and inequities in health care.<\/p>\n<p>Biological anthropologists now play a critical role in discrediting myths about race. By studying\u00a0 human variation, we can begin to understand evolutionary processes, adaptation, and the social implications of differences among human populations. Modern research shows that that humans are far more genetically homogenous than many other species, reinforcing a conclusion that \u2018race\u2019 is not a biological reality but a powerful social construct with real effects. Through evolutionary history and genetics, human diversity can be researched and understood, rather than than through racial categories.<\/p>\n<h2 class=\"import-Normal\">Review Questions<\/h2>\n<ul>\n<li class=\"import-Normal\">How is the genetic variation of the human species distributed worldwide?<\/li>\n<li class=\"import-Normal\">What evolutionary processes are responsible for producing genotypic\/phenotypic variation within and between human populations?<\/li>\n<li class=\"import-Normal\">Should we continue to attribute any value to \u201crace\u201d concepts older than 1950, based on our current understandings of human biological variation?<\/li>\n<li class=\"import-Normal\">How should we communicate scientific findings about human biological variation more accurately and responsibly to those outside the anthropological discipline?<\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\">Key Terms<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\"><strong>Age of Discovery<\/strong>: A period between the late 1400s and late 1700s when European explorers and ships sailed extensively across the globe in pursuit of new trading routes and territorial conquest.<\/p>\n<p class=\"import-Normal\"><strong>Ancestry<\/strong>: Biogeographical information about an individual, traced either through the study of an individual\u2019s genome, skeletal characteristics, or some other form of forensic\/archaeological evidence. Anthropologists carry out probabilistic estimates of ancestry. They attribute sets of human remains to distinctive \u201cancestral\u201d groups using careful statistical testing and should report ancestry estimations with statistical probability values.<\/p>\n<p class=\"import-Normal\"><strong>Binomial nomenclature<\/strong>: A system of naming living things developed by Linnaeus in the 1700s. It employs a scientific name made up of two italicized Latin or Greek words, with the first word capitalized and representative of an organism\u2019s genus and the second word indicating an organism\u2019s species (e.g., <em>Homo sapiens<\/em>, <em>Australopithecus afarensis<\/em>, <em>Pongo tapanuliensis<\/em>, etc.).<\/p>\n<p class=\"import-Normal\"><strong>Biological anthropology<\/strong>: A branch of study under anthropology (the study of humankind) that focuses on when and where humans and our human ancestors first originated, how we have evolved and adapted globally over time, and the reasons why we see biological variation among humans worldwide today.<\/p>\n<p class=\"import-Normal\"><strong>Biological determinism<\/strong>: The erroneous concept that an individual\u2019s behavioural characteristics are innate and determined by genes, brain size, or other physiological attributes\u2014and, notably, without the influence of social learning or the environment around the individual during development.<\/p>\n<p class=\"import-Normal\"><strong>Bony labyrinth<\/strong>: A system of interconnected canals within the auditory (ear- or hearing-related) apparatus, located in the inner ear and responsible for balance and the reception of sound waves.<\/p>\n<p class=\"import-Normal\"><strong>Cline<\/strong>: A gradient of physiological or morphological change in a single character or allele frequency among a group of species across environmental or geographical lines (e.g., skin colour varies clinally, as, over many generations, human groups living nearer the equator have adapted to have more skin pigmentation).<\/p>\n<p class=\"import-Normal\"><strong>Continuous variation<\/strong>: This term refers to variation that exists between individuals and cannot be measured using distinct categories. Instead, differences between individuals within a population in relation to one particular trait are measurable along a smooth, continuous gradient.<\/p>\n<p class=\"import-Normal\"><strong>Cystic fibrosis<\/strong>: A genetic disorder in which one defective gene causes overproduction and buildup of mucus in the lungs and other bodily organs. It is most common in northern Europeans (but also occurs in other world populations).<\/p>\n<p class=\"import-Normal\"><strong>Ecological niche<\/strong>: The position or status of an organism within its community and\/or ecosystem, resulting from the organism\u2019s structural and functional adaptations (e.g., bipedalism, omnivorous diets, lactose digestion, etc.).<\/p>\n<p class=\"import-Normal\"><strong>Essentialism<\/strong>: A belief or view that an entity, organism, or human grouping has a specific set of characteristics that are fundamentally necessary to its being and classification into definitive categories.<\/p>\n<p class=\"import-Normal\"><strong>Ethnicity<\/strong>: A term used commonly in an interchangeable way with the term <em>race<\/em>, complicated because how different people define this term depends on the qualities and characteristics they use to assign a label or identity to themselves and\/or others (which may include aspects of family background, skin colour, language(s) spoken, religion, physical proportions, behaviour and temperament, etc.).<\/p>\n<p class=\"import-Normal\"><strong>Eugenics<\/strong>: A set of beliefs and practices that involves the controlled selective breeding of human populations with the hope of improving their heritable qualities, especially through surgical procedures like sterilization and legal rulings that affect marriage rights for interracial couples.<\/p>\n<p class=\"import-Normal\"><strong>Gene flow<\/strong>: A neutral (or nonselective) evolutionary process that occurs when genes get shared between populations.<\/p>\n<p class=\"import-Normal\"><strong>Genetic drift<\/strong>: A neutral evolutionary process in which allele frequencies change from generation to generation due to random chance.<\/p>\n<p class=\"import-Normal\"><strong>Heterogeneity<\/strong>: The quality of being diverse genetically.<\/p>\n<p class=\"import-Normal\"><strong>Homog<\/strong><strong>enous<\/strong>: The quality of being uniform genetically.<\/p>\n<p class=\"import-Normal\"><strong>Human diversity<\/strong>: Human diversity is a measure of variation that may describe how many different forms of human there are, separated or clustered into groups according to some genetic, phenotypic, or cultural trait(s). The term can be applied to culture (in which case humans can be described as significantly diverse) or genetics (in which case humans are not diverse because all humans on Earth share a majority of their genes).<\/p>\n<p class=\"import-Normal\"><strong>Human variation<\/strong>: Differences in biology, physiology, body chemistry, behaviour, and culture. By measuring these differences, we understand the degrees of variation between individuals, groups, populations, or species.<\/p>\n<p class=\"import-Normal\"><strong>Isolation-by-distance model<\/strong>: A model that predicts a positive relationship between genetic distances and geographical distances between pairs of populations.<\/p>\n<p class=\"import-Normal\"><strong>Monogenetic<\/strong>: Pertaining to the idea that the origin of a species is situated in one geographic region or time (as opposed to <em>polygenetic<\/em>).<\/p>\n<p class=\"import-Normal\"><strong>Mutation<\/strong>: A gene alteration in the DNA sequence of an organism. As a random, neutral evolutionary process that occurs over the course of meiosis and early cell development, gene mutations are possible sources of variation in any given human gene pool. Genetic mutations that occur in more than 1% of a population are termed <em>polymorphisms<\/em>.<\/p>\n<p class=\"import-Normal\"><strong>Natural selection<\/strong>: An evolutionary process whereby certain traits are perpetuated through successive generations, likely owing to the advantages they give organisms in terms of chances of survival and\/or reproduction.<\/p>\n<p class=\"import-Normal\"><strong>Nonconcordance<\/strong>: The fact of genes or traits not varying with one another and instead being inherited independently.<\/p>\n<p class=\"import-Normal\"><strong>Otherness<\/strong>: In postcolonial anthropology, we now understand \u201cothering\u201d to mean any action by someone or some group that establishes a division between \u201cus\u201d and \u201cthem\u201d in relation to other individuals or populations. This could be based on linguistic or cultural differences, and it has largely been based on external characteristics throughout history.<\/p>\n<p class=\"import-Normal\"><strong>Out-of-Africa model<\/strong>: A model that suggests that all humans originate from one single group of <em>Homo sapiens<\/em> in (sub-Saharan) Africa who lived between 100,000 and 315,000 years ago and who subsequently diverged and migrated to other regions across the globe.<\/p>\n<p class=\"import-Normal\"><strong>Physical anthropology<\/strong>: This used to be the more common name given to the subdiscipline of anthropology centred upon the study of human origins, evolution and variation (also see <em>biological anthropology<\/em> above). This name for the field has gradually become less popular due to two reasons: first, it may not reflect our interests in other aspects of humankind that are not physical (such as those behavioural, cultural and spiritual), and second, using this term popular in the early decades of our field may be viewed by some as harkening back to a time when biological anthropologists conducted their work in unethical ways.<\/p>\n<p class=\"import-Normal\"><strong>Polygenetic<\/strong>: Having many different ancestries, as in older theories about human origins that involved multiple traditional groupings of humans evolving concurrently in different parts of the world before they merged into one species through interbreeding and\/or intergroup warfare. These earlier suggestions have now been overwhelmed by insurmountable evidence for a single origin of the human species in Africa (see the \u201cOut-of-Africa model\u201d).<\/p>\n<p class=\"import-Normal\"><strong>Polymorphism<\/strong>: A genetic variant within a population (caused either by a single gene or multiple genes) that occurs at a rate of over 1% among the population. Polymorphisms are responsible for variation in phenotypic traits such as blood type and skin colour.<\/p>\n<p class=\"import-Normal\"><strong>Population<\/strong>: A group of humans living in a particular geographical area, with more local interbreeding within-group than interbreeding with other groups. A limited or restricted amount of gene flow between populations can occur due to geographical, cultural, linguistic, or environmental factors.<\/p>\n<p class=\"import-Normal\"><strong>Population bottlenecking<\/strong>: An event in which genetic variation is significantly reduced owing to a sharp reduction in population size. This can occur when environmental disaster strikes or as a result of human activities (e.g., genocides or group migrations). An important example of this loss in genetic variation occurred over the first human migrations out of Africa and into other continental regions.<\/p>\n<p class=\"import-Normal\"><strong>Prejudice<\/strong>: An unjustified attitude toward an individual or group that is not based on reason, whether positive (and showing preference for one group of people over another) or negative (and resulting in harm or injury to others).<\/p>\n<p class=\"import-Normal\"><strong>Race<\/strong>: The identification of a group based on a perceived distinctiveness that makes that group more similar to each other than they are to others outside the group. This may be based on cultural differences, genetic parentage, physical characteristics, behavioural attributes, or something arbitrarily and socially constructed. As a social or demographic category, perceptions of \u201crace\u201d can have real and serious consequences for different groups of people. This is despite the fact that biological anthropologists and geneticists have demonstrated that all humans are genetically homogenous and that more differences can be found within populations than between them in the overall apportionment of human biological variation. This term is sometimes used interchangeably with <em>ethnicity<\/em>.<\/p>\n<p class=\"import-Normal\"><strong>Racism<\/strong>: Any action or belief that discriminates against someone based on perceived differences in race or ethnicity.<\/p>\n<p class=\"import-Normal\"><strong>Scientific Revolution<\/strong>: A period between the 1400s and 1600s when substantial shifts occurred in the social, technological, and philosophical sense, when a scientific method based on the collection of empirical evidence through experimentation was emphasized and inductive reasoning was used to test hypotheses and interpret their results.<\/p>\n<p class=\"import-Normal\"><strong>Typolog<\/strong><strong>ical<\/strong>: Of or describing an assortment system that relies on the interpretation of qualitative similarities or differences in the study of variation among objects or people. The categorization of cultures or human groups according to \u201crace\u201d was performed with a typological approach in the earliest practice of anthropology, but this practice has since been discredited and abandoned.<\/p>\n<p class=\"import-Normal\"><strong>Variance<\/strong>: In statistics, variance measures the dispersal of a set of data around the mean or average value.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration<strong><br \/>\n<\/strong><\/h2>\n<h3 class=\"import-Normal\"><strong>Videos<\/strong><\/h3>\n<p>American Medical Association (AMA). 2020. \u201c<a href=\"https:\/\/www.youtube.com\/watch?v=tqA3KvvscYc\" target=\"_blank\" rel=\"noopener\">Examining Race-Based Medicine<\/a>.\u201d YouTube, October 29. Accessed June 4, 2023.<\/p>\n<p>Crenshaw, Kimberl\u00e9. 2016. \u201c<a href=\"https:\/\/www.youtube.com\/watch?v=akOe5-UsQ2o\" target=\"_blank\" rel=\"noopener\">The Urgency of Intersectionality<\/a>.\u201d YouTube, December 7. Accessed June 4, 2023.<\/p>\n<p>Golash-Boza, Tanya. 2018. \u201c<a href=\"https:\/\/www.youtube.com\/watch?v=NQOimokvJXo\" target=\"_blank\" rel=\"noopener\">What Is Race? What Is Ethnicity? Is There a Difference?<\/a>.\u201d YouTube, October 28. Accessed June 4, 2023.<\/p>\n<p>Lasisi, Tina. 2020. \u201c<a href=\"https:\/\/naturalhistory.si.edu\/education\/teaching-resources\/social-studies\/webinar-how-hair-reveals-futility-race-categories\" target=\"_blank\" rel=\"noopener\">How Hair Reveals the Futility of Race Categories<\/a>.\u201d National Museum of Natural History webinar, October 21.<\/p>\n<p>Lasisi, Tina. 2022. \u201c<a href=\"https:\/\/www.youtube.com\/watch?v=_BEJvVFxKV4\" target=\"_blank\" rel=\"noopener\">Where Does My Skin Color Come From?<\/a>.\u201d PBS Terra, August 18. Accessed June 4, 2023.<\/p>\n<p>PBS Origins. 2018. \u201c<a href=\"https:\/\/www.youtube.com\/watch?v=CVxAlmAPHec\" target=\"_blank\" rel=\"noopener\">The Origin of Race in the USA<\/a>.\u201d YouTube, April 3. Accessed June 4, 2023.<\/p>\n<p>Roberts, Dorothy. 2016. \u201c<a href=\"https:\/\/www.youtube.com\/watch?v=KxLMjn4WPBY\" target=\"_blank\" rel=\"noopener\">The Problem with Race-Based Medicine<\/a>.\u201d YouTube, March 4. Accessed June 4, 2023.<\/p>\n<p>Vox. 2015. \u201c<a href=\"https:\/\/www.youtube.com\/watch?v=VnfKgffCZ7U\" target=\"_blank\" rel=\"noopener\">The Myth of Race, Debunked in 3 Minutes<\/a>.\u201d YouTube, January 13. Accessed June 4, 2023.<\/p>\n<h3 class=\"import-Normal\"><strong>Podcast Episodes<\/strong><\/h3>\n<p>Kwong, Emily, and Rebecca Ramirez. 2021. \u201c<a href=\"https:\/\/www.npr.org\/2021\/10\/05\/1043391809\/heres-a-better-way-to-talk-about-hair\" target=\"_blank\" rel=\"noopener\">Here\u2019s a Better Way to Talk about Hair: A 16 Minute Listen with Tina, Lasisi<\/a>\u201d NPR Short Wave, October 6. Accessed June 4, 2023.<\/p>\n<p>Speaking of Race. 2020. \u201c<a href=\"https:\/\/soundcloud.com\/user-88955638\/sets\/race-and-health?fbclid=IwAR2U1jdQL3XYFS5llGvYZ6uSrvPikuakmbxUZb--8voxgAMKrLbu7Ym7LGU\" target=\"_blank\" rel=\"noopener\">Race and Health series<\/a>.\u201d Speaking of Race, April 10. Accessed June 4, 2023.<\/p>\n<h3 class=\"import-Normal\"><strong>Websites<\/strong><\/h3>\n<p>Choices Program. 2023. \u201c<a href=\"https:\/\/www.choices.edu\/teaching-news-lesson\/an-interactive-timeline-black-activism-and-the-long-fight-for-racial-justice\/\" target=\"_blank\" rel=\"noopener\">An Interactive Timeline: Black Activism and the Long Fight for Racial Justice<\/a>.\u201d <em>Choices Program, Brown University<\/em> [Interactive Timeline], Updated February, 2023.<\/p>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">American Association of Biological Anthropologists. 2020. \u201c<a href=\"https:\/\/bioanth.org\/about\/position-statements\/open-letter-our-community-response-police-brutality-against-african-americans-and-call-antiracist-action\/\" target=\"_blank\" rel=\"noopener\">An Open Letter to Our Community in Response to Police Brutality against African-Americans and a Call to Antiracist Action<\/a>\u201d. <em>American Association of Biological Anthropologists<\/em>, June 10, 2020. 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Girgis, et al. 1984. \u201cBrain Size, Cranial Morphology, Climate, and Time Machines [and Comments and Reply].\u201d <em>Current Anthropology<\/em> 25 (3): 301\u2012330.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Betti, Lia, Fran\u00e7ois Balloux, Tsunehiko Hanihara, and Andrea Manica. 2010. \u201cThe Relative Role of Drift and Selection in Shaping the Human Skull.\u201d <em>American Journal of Physical Anthropology<\/em> 141 (1): 76\u201282. https:\/\/doi.org\/10.1002\/ajpa.21115.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Betti, Lia, Fran\u00e7ois Balloux, William Amos, Tsunehiko Hanihara, and Andrea Manica. 2009. \u201cDistance from Africa, Not Climate, Explains Within-Population Phenotypic Diversity in Humans.\u201d <em>Proceedings: Biological Sciences<\/em> 276 (1658): 809\u2012814. https:\/\/doi.org\/10.1098\/rspb.2008.1563.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Betti, Lia, Noreen von Cramon-Taubadel, Andrea Manica, and Stephen J. 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Feldman. 2002. \u201cGenetic Structure of Human Populations.\u201d <em>Science<\/em> 298 (5602): 2381\u20122385.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Sauer, Norman J. 1992. \u201cForensic Anthropology and the Concept of Race: If Races Don\u2019t Exist, Why Are Forensic Anthropologists So Good at Identifying Them?\u201d <em>Social Science and Medicine<\/em> 34 (2): 107\u2012111. https:\/\/doi.org\/10.1016\/0277-9536(92)90086-6.<\/p>\n<p class=\"import-Normal\">Shonkoff, Jack P., Natalie Slopen, and David R. Williams. 2021. \u201cEarly Childhood Adversity, Toxic Stress, and the Impacts of Racism on the Foundations of Health.\u201d <em>Annual Review of Public Health<\/em> 42: 115\u2012134.<\/p>\n<p class=\"import-Normal\">Sjoding, Michael W., Robert P. Dickson, Theodore J. Iwashyna, Steven E. Gay, and Thomas S. Valley. 2020. \u201cRacial Bias in Pulse Oximetry Measurement.\u201d <em>The New England Journal of Medicine<\/em> 383: 2477-2478.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Staes, Nicky, Chet C. Sherwood, Katharine Wright, Marc de Manuel, Elaine E. Guevara, Tomas Marques-Bonet, Michael Kr\u00fctzen, et al. 2017. \u201cFOXP2 Variation in Great Ape Populations Offers Insight into the Evolution of Communication Skills.\u201d <em>Scientific Reports<\/em> 7 (1): 1\u201210. https:\/\/doi.org\/10.1038\/s41598-017-16844-x.<\/p>\n<p class=\"import-Normal\">Tomati, Opal. 2016. \u201cAn Interview with the Founders of Black Lives Matter.\u201d TED Talks 2016, October 26\u201228. Accessed June 15, 2023. https:\/\/www.ted.com\/talks\/alicia_garza_patrisse_cullors_and_opal_tometi_an_interview_with_the_founders_of_black_lives_matter\/up-next.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Tsai, Jennifer. 2021. \u201cCOVID-19 Is Not a Story of Race, but a Record of Racism\u2014Our Scholarship Should Reflect That Reality.\u201d <em>The American Journal of Bioethics<\/em> 21 (2): 43\u201247. https:\/\/doi.org\/10.1080\/15265161.2020.1861377.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">von Cramon-Taubadel, Noreen, and Stephen J. Lycett. 2008. \u201cBrief Communication: Human Cranial Variation Fits Iterative Founder Effect Model with African Origin.\u201d <em>American Journal of Physical Anthropology<\/em> 136 (1): 108\u2012113. https:\/\/doi.org\/10.1002\/ajpa.20775.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Weiss, Kenneth M., and Jeffrey C. Long. 2009. \u201cNon-Darwinian Estimation: My Ancestors, My Genes\u2019 Ancestors.\u201d <em>Genome Research<\/em> 19: 703\u2012710. https:\/\/doi.org\/10.1101\/gr.076539.108.19.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Williams, David W. 2018. \u201cStress and the Mental Health of Populations of Color: Advancing Our Understanding of Race-related Stressors.\u201d <em>Journal of Health and Social Behavior<\/em> 59 (4): 466\u2012485.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Wise, Tim. 2010. <em>Colorblind: The Rise of Post-Racial Politics and the Retreat from Racial Equity<\/em>. San Francisco: City Lights.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Yudell, Michael, Dorothy Roberts, Rob DeSalle, and Sarah Tishkoff. 2016. \u201cTaking Race out of Human Genetics.\u201d <em>Science<\/em> 351 (6273): 564\u2012565. https:\/\/doi.org\/10.1126\/science.aac4951.<\/p>\n<\/div>\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_1683_1696\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1696\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Leslie E. Fitzpatrick, Ph.D., Independent Archaeological Consultants<\/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__-13\/\"><em>Chapter 14: Human Variation: An Adaptive Significance Approach<\/em><\/a><em>\u201d by Leslie E. Fitzpatrick. 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\">Learning Objectives<\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Distinguish between adaptations and adjustments as ways of coping with environmental stressors.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Provide examples of adjustments humans use to cope with thermal stressors.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Describe how specific patterns of human adaptations and adjustments are correlated to natural selection processes.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Summarize the role of solar radiation in variations of human skin tone, and explain why reduced pigmentation is advantageous in northern latitudes.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Compare and contrast the various genetic mutations present in Tibetan and Ethiopian populations that allow them to survive at high altitudes.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Define the relationship between specific genetic mutations in some human populations and certain infectious diseases, such as the sickle-cell trait mutation and malarial infection.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p>As early humans left Africa and spread across the globe, they faced numerous challenges related to their new environments. Beyond genetically influenced changes in physiology as a result of evolution, humans have developed lifestyle strategies to cope with and even thrive in a wide range of habitats. The ways populations of humans met such challenges, coupled with their geographic separation throughout the majority of the last two hundred thousand years, have led to the many forms of adaptation in our species. This chapter focuses on the complexities of modern human variation through the lens of human evolutionary history.<\/p>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Stress and Homeostasis<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">All organisms, including humans, must maintain a baseline of normal functions within their cells, tissues, and organs to survive. This constancy of internal functions is referred to as <strong>homeostasis.<\/strong> Homeostatic regulation, however, may be disrupted by a variety of both external and internal stimuli known as <strong>stressors<\/strong>. Within limits, all organisms have evolved certain physiological mechanisms to respond to stressors in an effort to maintain homeostasis. The range of changes in the physiology (function), morphology (form), and\/or behaviour of organisms in response to their environments and potential stressors is regulated by its <strong>phenotypic<\/strong><strong> plasticity<\/strong>. Coping with these stressors led to the development of both <strong>adjustments<\/strong> (behavioural, acclimatory, and developmental) and <strong>adaptations<\/strong>, which are explained in detail in the following sections.<\/p>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Adjustments and Adaptations<\/h2>\n<h3 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Adjustments<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The term <em>adjustment<\/em> refers to an organism\u2019s nongenetic way of coping with the stressors of its environment. Although adjustments themselves are nongenetic in nature, the ability of an organism to experience or develop an adjustment is based on its phenotypic plasticity, which is linked to its evolutionarily guided genetic potential. Adjustments occur exclusively on the individual level. As such, different individuals within a population may experience a wide range of possible adjustments in response to a similar stressor. In general, the three main forms of adjustment are: behavioural, acclimatory, and developmental.<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><em>Behavioural Adjustments<\/em><\/h4>\n<figure style=\"width: 390px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2023\/06\/image3-3.jpg\" alt=\"Underwater photograph of scuba diver exploring the ocean floor.\" width=\"390\" height=\"293\" \/><figcaption class=\"wp-caption-text\">Figure 15.1: Notice the lack of full-spectrum colour in this photo of a deep-water diver as well as the diver\u2019s use of specialized equipment, such as a breathing apparatus to deliver gasses for respiration, a bodysuit to ensure thermal regulation, and a flashlight to increase visibility in the low-light setting. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-13\/\">Deep water diver<\/a> by Leslie E. Fitzpatrick 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\">When you are cold, do you reach for a blanket? When you are warm, do you seek out shelter cooled by an air-conditioning system? If so, you have likely been influenced to do so by the culture in which you were raised. As noted earlier in this textbook, the term <em>culture<\/em> refers to a collection of shared, learned beliefs and behaviours among individuals within a discrete population. <strong>Behavioural adjustments<\/strong> are regarded as cultural responses to environmental stressors. These adjustments are temporary in nature and, since they are nongenetic, must be constantly altered to meet novel situations posed by the environment. For example, divers are able to reach extraordinary depths (in excess of 300 metres below the surface) within the water through the use of a specialized mixture of gasses for breathing, an apparatus for the delivery of the gasses, protective clothing, and gear to increase visibility. The deeper a diver descends, the more atmospheric pressure the diver experiences, resulting in increased levels of potentially toxic byproducts of respiration within the body. In addition, with increased depth there is a decrease in the ambient temperature of the water and a decrease in the availability of light within the visible spectrum. Deep-water divers are well-versed in the environmental stressors of open waters and employ a variety of strategies based on behavioural adjustments to meet such demands. From wearing protective clothing to help maintain the body\u2019s core temperature to waiting at a specific depth for a prescribed period of time to facilitate the expulsion of nitrogen gas that may have accumulated within the bloodstream, divers employ numerous behavioural adjustments to ensure their safety (Figure 15.1). Without these culturally mediated behavioural adjustments, a deep-water diver\u2019s first dive would also be their last.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In many developing countries, the use of refrigeration for the storage of perishable food products is uncommon; therefore, individuals within these cultures have developed a variety of behavioural adjustment strategies related to food preparation to address possible food spoilage. Through a cross-cultural analysis of spice use in recipes, Paul Sherman and Jennifer Billing (1999) determined that cultures closest to the equator, where temperatures are hotter, tend to use both a greater number and a wider variety of plant-based spices with bacteria-inhibiting phytochemical properties (e.g., garlic and onion). Antimicrobial properties of spices permits the consumption of foods, particularly animal-based protein sources, for a period of time beyond that which would be considered safe. There are some acclimatory adjustment benefits to the use of some pungent spices as well, which are explored in the following section.<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><em>Acclimatory Adjustments: Thermal Stressors<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Acclimatory adjustments<\/strong> are temporary, reversible changes in an organism\u2019s physiology in response to environmental stressors. Although they are not genetically determined, the range of acclimatory adjustments that an organism is capable of producing is linked to its underlying phenotypic plasticity and the duration and severity of the stressor. A good example of this is the human response to varying ambient temperatures.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">To understand human adjustments, we must first understand the thermodynamic mechanisms through which heat may be gained or lost. The four pathways for this are conduction, convection, evaporation, and radiation (Figure 15.2).<\/p>\n<figure style=\"width: 376px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image11-5.png\" alt=\"Person facing wall with arrows illustrating evaporation, conduction, radiation, and convection.\" width=\"376\" height=\"426\" \/><figcaption class=\"wp-caption-text\">Figure 15.2: Various thermodynamic mechanisms related to heat gain and loss in the human body. This process is decribed in the text below. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-13\/\">Mechanisms of heat transfer (Figure 14.2)<\/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 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Through <strong>conduction<\/strong> processes, heat will move from a warmer body to a cooler one through direct contact. An example of this is when you accidentally touch a hot cooktop with your hand and the heat is transferred from the cooktop to your skin.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">With <strong>convection<\/strong>, when a warm body is surrounded by a cooler fluid (e.g., air or water), heat will be transferred from the warmer body to the cooler fluid. This is why we will often employ the behavioural adjustment of wearing multiple layers of clothing during the winter in an effort to prevent heat loss to the cooler atmosphere. Conversely, if your body temperature is cooler than that of the air surrounding you, your body will absorb heat.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Depending on your physical condition, most people will begin to sweat around 37.2\u2103 to 37.7\u2103 (98.9\u2109\u201399.9\u2109). Sweating is an example of <strong>evaporation<\/strong>, which occurs when a liquid, such as the water within our bodies, is converted to a gas. Phase conversions, such as those underlying the evaporative processes of transforming liquids to gasses, require energy. In evaporation, this energy is in the form of heat, and the effect is to cool the body.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The final mechanism for heat loss within the human body is <strong>radiation<\/strong>, through which energy in the form of electromagnetic waves is produced at a wavelength that typically lies below that which is visible to the human eye. Although humans gain and lose heat from their bodies through radiation, this form of heat transfer is not visible. Humans are capable of losing and gaining heat through conduction, convection, and radiation; however, heat may not be gained through evaporation.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">As the ambient temperature decreases, it becomes increasingly difficult for the human body to regulate its core temperature, which is central to the maintenance of homeostasis. When an individual\u2019s body temperature falls below 34.4\u2103 (93.9\u00b0F), the brain\u2019s <strong>hypothalamus<\/strong> becomes impaired, leading to issues with body temperature control. A total loss of the ability to regulate body temperature occurs around 29.4\u2103 (84.9\u00b0F), which may result in death. When the ambient temperature falls below the critical temperature of 31\u2103 (87.8\u00b0F), a nude human body that is at rest will respond with a series of physiological changes to preserve homeostasis (Figure 15.3).<\/p>\n<figure style=\"width: 585px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image26-2.jpg\" alt=\"Body outlines illustrate differences in heat maintenance for different ambient environments.\" width=\"585\" height=\"347\" \/><figcaption class=\"wp-caption-text\">Figure 15.3: Example of overall body heat maintenance in cold and warm ambient environments. <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__-13\/\">Body heat maintenance in cold and warm (Figure 14.3)<\/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=\"margin-left: 0pt;text-indent: 0pt\">The human body experiences two main types of physiological responses to colder temperatures: those that increase the production of heat and those that seek to retain heat. The production of heat within the body is accomplished through short-term increases in the body\u2019s basal metabolic rate, such as shivering to increase muscular metabolism. An organism\u2019s basic metabolic rate is a measure of the energy required to maintain necessary body processes when the organism is at rest. Increases in basal metabolic rates, such as when we shiver from the cold, require increased consumption of energy-providing nutrients. Of course, such increases in metabolic rates are not infinite, as we may only consume a finite amount of nutrients. As with all <strong>acclimatory adjustments<\/strong>, an increase in the basal metabolic rate is merely temporary.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Of the physiological mechanisms to preserve heat already in the body, the most notable is <strong>vasoconstriction<\/strong>, or the constriction of peripheral capillaries in the skin. The decreased surface area of the capillaries through vasoconstriction results in less heat reaching the surface of the skin where it would be dissipated into the atmosphere. Vasoconstriction also leads to the maintenance of heat near the core of the body where the vital organs are located. As a trade-off, though, individuals are more at risk of cold-related injuries, such as frost-bite, which can lead to tissue necrosis (tissue death) in regions of the body that are most distant from the core (e.g., fingers, toes, nose, ears, cheeks, chin, etc.).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Just as cold stress presents challenges to maintaining homeostasis, heat does as well. In hot climates, the body will absorb heat from its surroundings (through conduction, convection, and radiation), resulting in potential heat-related disorders, such as heat exhaustion. When the human body is exposed to ambient temperatures above 35\u2103 (95\u00b0F), excess body heat will be lost primarily through evaporative processes, specifically through sweating. All humans, regardless of their environment, have approximately the same number of sweat glands within their bodies. Over time, individuals living in hot, arid environments will develop more sensitive forms of sweat glands resulting in the production of greater quantities of sweat (Best, Lieberman, and Kamilar 2019; Pontzer Et al. 2021). In an effort to prevent dehydration due to this form of acclimatory adjustment, there will be an additional reduction in the volume of urine produced by the individual (Pontzer Et al. 2021).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">As noted in the previous section, some cultural groups, particularly those in equatorial regions, add pungent spices to their foods to inhibit the colonization of bacteria (Sherman and Billing 1999). Although adding spices to decrease spoilage rates is a behavioural adjustment, the application of some forms of peppers triggers an acclimatory adjustment as well. Compounds referred to as capsaicinoids are the secondary byproducts of chili pepper plants\u2019 metabolism and are produced to deter their consumption by some forms of fungi and mammals. When mammals, such as humans, consume the capsaicinoids from chili peppers, a burning sensation may occur within their mouths and along their digestive tracts. This burning sensation is the result of the activation of capsaicin receptors along the body\u2019s nerve pathways. Although the peppers themselves may be at ambient temperature so their consumption is not causing any form of body temperature increase, the human body perceives the pepper as elevating its core temperature due to the activation of the capsaicin receptors. This causes the hypothalamus to react, initiating sweating in an attempt to lower body temperature and maintain homeostasis. The increased piquancy (application of pungent spices to food) as a means of inhibiting food-borne bacterial colonization in warm climates, as well as spices\u2019 ability to trigger sweating processes as a method for cooling the body, is an example of the intersection between behavioural and acclimatory adjustments that utilized within certain populations.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In addition to increased sweat production to maintain homeostasis in excess heat, <strong>vasodilation<\/strong> may occur (Figure 15.4). Vasodilation is an expansion of the capillaries within the skin leading to a more effective transfer of heat from within the body to the exterior to allow conductive, convective, radiative, and evaporative (sweating) processes to occur.<\/p>\n<figure style=\"width: 499px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image27-4.png\" alt=\"Side-by-side sketches of capillaries under the skin for heat and cold.\" width=\"499\" height=\"445\" \/><figcaption class=\"wp-caption-text\">Figure 15.4: The vasoconstriction processes occur within the peripheral vascular system when an individual is exposed to cold ambient temperatures and the vasodilation that occurs in warmer environments. <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__-13\/\">Vasoconstriction and vasodilation (Figure 14.4)<\/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\">Physiologically based acclimatory adjustments to hot, dry climates may be complemented by behavioural adjustments as well. For example, individuals in such climates may limit their physical activity during the times of day when the temperature is typically the hottest. Additionally, these individuals may wear loose-fitting clothing that covers much of their skin. The looseness of the clothing allows for air to flow between the clothing and the skin to permit the effective evaporation of sweat. Although it may seem counterintuitive to cover one\u2019s body completely in a hot climate, the covering of the skin keeps the sun\u2019s rays from directly penetrating the skin and elevating the body\u2019s core temperature.<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><em>Acclimatory Adjustments: Altitudinal Stressors<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The challenges posed by thermal conditions are but one form of environmental stressor humans must face. High-altitude environments, which are defined as altitudes in excess of 2,400 metres above sea level (masl) or 7,874 feet above sea level (fasl), pose additional challenges to the maintenance of homeostasis in humans. Some of the main stressors encountered by those living within high-altitude environments include decreased oxygen availability, cold temperatures, low humidity, high wind speed, a reduced nutritional base, and increased solar radiation levels. Of these challenges, the most significant is the decreased availability of oxygen.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">To visualize how altitude affects the availability of oxygen, imagine two balloons that are each filled with the same quantity of oxygen molecules. One of these balloons is positioned at sea-level and the other is placed high upon a mountain peak. For the balloon at sea level, there is more atmospheric pressure pressing down on the molecules within this balloon. This leads to the oxygen molecules within the sea level balloon being forced into a more compact organization. In contrast, the mountain peak balloon has less atmospheric pressure pressing down on it. This leads to the oxygen molecules within that balloon spreading out from each other since they are not being forced together quite as strongly. This example highlights the availability of oxygen molecules in each breath than we take in low- versus high-altitude environments. At 5,500 masl (approximately 18,000 fasl), the atmospheric pressure is approximately 50% of its value at sea level (Peacock 1998). At the peak of Mount Everest (8,900 masl or approximately 29,200 fasl), the atmospheric pressure is equivalent to only about 30% of their sea level amounts (Peacock 1998; Figure 15.5).<\/p>\n<figure style=\"width: 482px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image21-5.png\" alt=\"Atmospheric pressure decreases in density as a person climbs a mountain and increases in altitude. \" width=\"482\" height=\"418\" \/><figcaption class=\"wp-caption-text\">Figure 15.5: As altitude increases, atmospheric pressure decreases, which allows for more space between air molecules. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-13\/\">Atmospheric pressure (Figure 14.5)<\/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\">Due to decreased availability of oxygen at higher altitudes, certain acclimatory adjustments are required to ensure the maintenance of homeostasis for individuals other than those who were gestated, born, and raised at high altitude. For these people, their rate of breathing will increase to permit greater quantities of air containing oxygen into the lungs when they ascend into higher altitude environments. An increased speed and depth of breathing, which is referred to as <strong>hyperpnea<\/strong>, is not sustainable indefinitely; thus, the rate of breathing begins to decrease as the person becomes acclimatized to the altitude. During the initial phases of high-altitude-related hyperpnea, the heart begins to beat faster but the amount of blood pushed through during each beat decreases slightly. In addition, the body will divert energy from noncritical bodily functions, such as digestive processes.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Once the atmospheric oxygen reaches the alveoli (small air sacs) in the lungs, it spreads across the alveolar membrane and enters <strong>erythrocytes<\/strong> (red blood cells). As oxygen reaches the alevoli\u2019s erythrocytes, it loosely binds with hemoglobin (an iron-rich protein) contained in the erythrocytes. When the erythrocytes carrying the hemoglobin-bound oxygen molecules reach capillaries where the partial pressure of oxygen is relatively low, oxygen will be released by the hemoglobin so that it is free for diffusion into body cells. Similar to acclimatory adjustments related to thermal conditions (e.g., shivering or sweating), those related to high altitude may not be infinitely sustained due to their energetically expensive nature.<\/p>\n<figure style=\"width: 430px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image24-3.jpg\" alt=\"Newborn lays face down on hospital bedding. Photograph is blue indicating UV light.\" width=\"430\" height=\"323\" \/><figcaption class=\"wp-caption-text\">Figure 15.6: Premature infant born at 30 weeks, 4 days gestation to a mother with altitudinal-induced preeclampsia. Blue light assists the infant\u2019s liver with processing high levels of bilirubin. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-13\/\">Premature infant<\/a> by Leslie E. Fitzpatrick 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\">Although the long-term acclimatory adjustments that an individual from low altitude experiences in a high-altitude environment may permit them to reside there successfully, reproduction within such settings is frequently complicated. With increased altitude comes an increased risk of miscarriage, lower birth weights, and higher infant mortality rates. As the pregnant person\u2019s body seeks to preserve its own homeostasis, there is often a decreased rate and volume of blood flow to the uterus as compared to a pregnant person of similar physiological condition at a lower altitude (Moore, Niermeyer, and Zamudio 1998). This results in a decrease in the amount of oxygen that will be passed through the uterus and placenta to the developing fetus. In addition, pregnant people who experience pregnancy at higher altitudes are more prone to developing preeclampsia (severe elevation of blood pressure), which is linked to increased rates of both fetal and maternal death (Moore, Niermeyer, and Zamudio 1998; Figure 15.6).<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><em>Developmental Adjustments <\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Developmental adjustments occur only in individuals who spent their developmental period (i.e., childhood and adolescence) within a high-altitude environment; they do not apply to those who moved into these environments in the post developmental (i.e., adult) phase. Furthermore, the degree of developmental adjustment within an individual is directly related to their underlying phenotypic plasticity as well as the amount of time during the crucial growth and development period that the individual resides within the challenging environment. Although humans have the remarkable capacity to develop and survive within environments that are not overly conducive to the successful maintenance of homeostasis, there are definitely physiological costs associated with this ability.<\/p>\n<figure style=\"width: 369px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image18-2.jpg\" alt=\"Two people walking down a street wearing brightly colored woven traditional clothing.\" width=\"369\" height=\"277\" \/><figcaption class=\"wp-caption-text\">Figure 15.7: Two individuals from a high-altitude region of the Peruvian Andes. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Andahuaylas_Peru-_two_women_walking_down_street.jpg\">Andahuaylas Peru-two women walking down street<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:ThayneT\">Thayne Tuason<\/a> has been modified (cropped) and is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/deed.en\">CC BY 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In general, high-altitude natives tend to grow more slowly and physically mature later than their low-altitude counterparts (Figure 15.7). Lowered growth and maturity rates are linked not only to the increased physiological demands placed on the body due to the decreased partial pressure of oxygen but also to reductions in the quality of the nutritional base at higher altitudes. Increased terrain complexity, elevated solar radiation levels, and higher wind speeds coupled with the lower temperatures and humidity levels found at high altitudes leads to difficulties with growing and maintaining crops and raising livestock. Overall, as altitude rises, the quality of the available nutritional base goes down, which is correlated to a lack of the nutrients necessary to ensure proper physiological growth and development in humans. Thus, even though individuals may be able to develop and grow within high-altitude environments, they may not reach their full genetically mediated growth potential as they would in a lower-altitude environment.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Not all developmental adjustments are linked to environmental pressures such as climate or altitude; rather, some of these adjustments are correlated to sociocultural or behavioural practices. Some of these adjustments may affect the physiological appearance of an individual when they are practiced consistently during the development and growth phases.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Sudden infant death syndrome (SIDS) has no definitive cause; however, the American Academy of Pediatrics published a report in 1992 linking SIDS to infants (under the age of one) sleeping on their stomachs. The \u201cBack to Sleep\u201d campaign championed by the American Academy of Pediatrics helped educate members of the medical community as well as the public that the best sleep position for infants is on their backs (American Academy of Pediatrics 2000).<\/p>\n<\/div>\n<figure id=\"attachment_420\" aria-describedby=\"caption-attachment-420\" style=\"width: 562px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-419\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/14.8.jpg\" alt=\"Brachycephaly drawings show a flattened, symmetrical head. Plagiocephaly shows asymmetrical flattening of the skull.\" width=\"562\" height=\"219\" \/><figcaption id=\"caption-attachment-420\" class=\"wp-caption-text\">Figure 15.8: These sketches illustrate a top and side view of brachycephaly (left and middle images, respectively) and plagiocephaly (right image). Credit: Brachycephaly and plagiocephaly original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) 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 class=\"__UNKNOWN__\">\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Placing infants on their backs to sleep has led to decreased infant mortality (death) rates due to SIDS; however, it has led to an unintended consequence: infant cranial deformation. The cranial deformations experienced by infants who sleep solely on their back tend to manifest in one of two forms: brachycephaly and plagiocephaly (Roby Et al. 2012; Figure 15.8). With positional brachycephaly, the back of the infant\u2019s head appears rather uniformly flattened due to repetitive contact with a flat surface, such as a crib mattress or car seat back. In cases of positional plagiocephaly, the back of the infant\u2019s head appears asymmetrically flattened. This asymmetry is typically due to an uneven distribution of mechanical forces resulting from the manner in which the infant\u2019s head is in contact with a flat surface. The forms of cranial deformation resulting from sleep positioning do not affect the infant\u2019s brain development. For many individuals, the appearance of the deformation is minimized during later development. Still, some individuals will maintain the pattern of cranial deformation acquired during their infancy throughout their lives. The unintentional cranial deformation resulting from placing infants on their backs to sleep as a means of preventing SIDS-related deaths is a physiological indicator of a behavioural adjustment.<\/p>\n<\/div>\n<h3 class=\"__UNKNOWN__\"><strong>Adaptations<\/strong><\/h3>\n<div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">As we have just explored, survival and reproduction at high altitudes present numerous physiological challenges for most humans. The behavioural, acclimatory, and developmental adjustments discussed above are all related to the phenotypic plasticity of the individual; however, most adjustments are temporary in nature and they affect a single individual rather than all individuals within a population. But what if the physiological changes were permanent? What if they affected all members of a population rather than just a single individual? The long-term, microevolutionary (i.e., genetic) changes that occur within a population in response to an environmental stressor are referred to as an adaptation. From an evolutionary standpoint, the term <em>adaptation<\/em> refers to a phenotypic trait (i.e., physiological\/morphological feature or behaviour) that has been acted upon by natural selection processes to increase a species\u2019 ability to survive and reproduce within a specific environment. Within the field of physiology, the term <em>adaptation<\/em> refers to traits that serve to restore homeostasis. The physiology-based interpretation of adaptations presumes that all traits serve a purpose and that all adaptations are beneficial in nature; however, this may be a fallacy, since some traits may be present without clear evidence as to their purpose. As such, during the following discussion of various forms of adaptations in human populations, we will focus our attention on phenotypic traits with an evidence-based purpose.<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><em>Adaptation: <\/em><em>Altitudinal Adaptation<\/em><\/h4>\n<\/div>\n<figure id=\"attachment_420-2\" aria-describedby=\"caption-attachment-420-2\" style=\"width: 619px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-420\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/14.9.jpg\" alt=\"The Simian Plateau in northeast Africa and Tibetan Plateau in southern Asia.\" width=\"619\" height=\"339\" \/><figcaption id=\"caption-attachment-420-2\" class=\"wp-caption-text\">Figure 15.9: Highlighted regions feature (from left to right) the Simian (Ethiopian) and Tibetan Plateau high-altitude regions. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:World_Map_of_HVR_adaptation_in_high_altitude_populations.jpg\">World Map of HVR adaptation in high altitude populations<\/a> by Chkuu has been modified (cropped, Andean region colour change) and is under <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<div class=\"__UNKNOWN__\">\n<p>As mentioned in the previous section, there is genomic research supporting the evolutionary selection of certain phenotypes and their corresponding genotypes within indigenous high-altitude populations across the globe. The following discussion focuses on two high-altitude indigenous populations from Tibet and Ethiopia (Figure 15.9). Although these populations share many common genetic traits based on relatively similar evolutionary histories influenced by similar environmental stressors, there is support for local genetically based adaptation as well, based on different genes being acted upon by environmental stressors that may be unique to Tibet and Ethiopia (Bigham 2016).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Tibetan populations have resided in the Tibetan Plateau and Himalayan Mountain regions at elevations exceeding 4,000 masl (13,100 fasl) for at least the past 7,400 years (Meyer Et al. 2017). There is evidence of a genetic exchange event involving Tibetan populations and Denisovans around 48,700 years ago, which introduced a haplogroup involving mutations of the <em>EPAS1<\/em> gene (Zhang Et al. 2021). The <em>EPAS1<\/em> is involved in the regulation of erythrocytes and hemoglobin. For individuals originating in lower-altitude environments, <em>EPAS1 <\/em>stimulates increased erythrocyte production in high-altitude environments as a temporary acclimatory adjustment. For indigenous high-altitude populations of Tibet, the <em>EPAS1<\/em> gene mutation introduced by Denisovan introgression inhibits increased erythrocyte production, which reduces potential negative effects (e.g., stroke or heart attack) associated with long-term high levels of erythrocyte production (Gray Et al. 2022; Zhang Et al. 2021). The erythrocyte count of high-altitude Tibetans with the <em>EPAS1<\/em> point mutation is about the same as for individuals residing at sea level.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Populations indigenous to the Semien Plateau of Ethiopia, such as the Oromo and Amhara, share a similar but not identical <em>EPAS1<\/em> point mutation with the Tibetan population (Bigham 2016); however, there is no indication that this mutation was derived from Denisovan introgression. The <em>EPAS1<\/em> mutations occurred independently from each other; however, their effects are still similar in that they permit the Tibetan and Ethiopian populations to survive at high altitudes. Not all adaptations are related to life in high-altitude environments, however. In the following sections, we will address two more general examples of adaptation in human populations: variations in skin colour and differences in body build.<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><em>Adaptation: Skin Tone<\/em><em> Basics<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">When you think about your own skin tone and compare it to members of your family, do you all possess exactly the same shade? Are some members of your family darker than others? What about your friends? Your classmates? Skin tone occurs along a continuum, which is a reflection of the complex evolutionary history of our species. The expression of skin tone is regulated primarily by melanin and hemoglobin. <strong>Melanin<\/strong> is a dark brown-black pigment that is produced by the oxidation of certain amino acids (e.g., tyrosine, cysteine, phenylalanine) in melanocytes. <strong>Melanocytes<\/strong> are specialized cells located in the base layer (stratum basale) of the skin\u2019s epidermis as well as several other areas within the body (Figure 15.10). Within the melanocytes, melanin is produced in the special organelle called a melanosome. Melanosomes serve as sites for the synthesis, storage, and transportation of melanin. Melanosomes transport the melanin particles through cellular projections to epidermal skin cells (keratinocytes) as well as to the base of the growing hair root. In the eye, however, melanin particles produced by the melanosomes remain present within the iris and are not transported beyond their origin location. The two main forms of melanin related to skin, hair, and eye colour are eumelanin and pheomelanin. All humans contain both eumelanin and pheomelanin within their bodies; however, the relative expression of these two forms of melanin determines an individual\u2019s overall colouring. Eumelanin is a brown-to-black coloured melanin particle while pheomelanin is more pink-to-red coloured. Individuals with darker skin or hair colour have a greater expression of eumelanin than those with lighter-coloured skin and blonde or red hair.<\/p>\n<figure style=\"width: 553px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image12-2.jpg\" alt=\"Melanocytes and melanosomes are compared from light and dark skin tones.\" width=\"553\" height=\"436\" \/><figcaption class=\"wp-caption-text\">Figure 15.10: Diagram featuring the relative numbers of melanocytes and melanosomes in light and dark shades of skin tone. <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:\/\/openstax.org\/books\/anatomy-and-physiology-2e\/pages\/5-1-layers-of-the-skin#fig-ch05_01_07\">Skin Pigmentation (Anatomy and Physiology, Figure 5.8)<\/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<h4 class=\"import-Normal\"><em>Adaptation<\/em><em>: <\/em><em>Melanogenesis <\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Although all humans have approximately the same number of melanocytes within their epidermis, the production of melanin by these melanocytes varies. There are two forms of melanogenesis (the process through which melanocytes generate melanin): basal and activated. As discussed previously, the expression of eumelanin and pheomelanin by the melanocytes is genetically regulated through the expression of specific receptors (e.g., <em>MC1R<\/em>) or other melanocyte components (e.g., <em>MFSD12<\/em>)<em>.<\/em> <strong>Basal melanogenesis <\/strong>is dependent upon an individual\u2019s inherent genetic composition and is not influenced by external factors. <strong>Activated melanogenesis<\/strong> occurs in response to ultraviolet radiation (UV) exposure, specifically UV-B (short UV wave) exposure. Increased melanogenesis in response to UV-B exposure serves to provide protection to the skin\u2019s innermost layer called the hypodermis, which lies below the epidermis and dermis (Figure 15.11). Melanin in the skin, specifically eumelanin, effectively absorbs UV-B radiation from light\u2014meaning that it will not reach the hypodermal layer. This effect is often more apparent during periods of the year when people tend to be outside more and the weather is warmer, which leads to most donning fewer protective garments. The exposure of skin to sunlight is, of course, culturally mediated with some cultures encouraging the covering of skin at all times.<\/p>\n<figure style=\"width: 395px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image10-6.png\" alt=\"Cross-section of skin illustrating UVA rays penetrate deeper than UVB. \" width=\"395\" height=\"407\" \/><figcaption class=\"wp-caption-text\">Figure 15.11: Penetration of skin layers by UVA and UVB rays. UVB rays penetrate only through the epidermis. UVA rays penetrate much deeper, through the dermis. Credit: <a href=\"https:\/\/cnx.org\/contents\/FPtK1zmh@16.7:RxywCGkA@10\/5-1-Layers-of-the-Skin\">Skin Pigmentation (Anatomy and Physiology, Figure 5.8)<\/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\">As previously noted in this chapter, hemoglobin is an iron-rich protein that binds with oxygen in the bloodstream. For individuals with lighter-coloured skin, blood vessels near the surface of the skin and the hemoglobin contained within those vessels is more apparent than in individuals with darker skin. The visible presence of hemoglobin coupled with the pink-to-red tone of the pheomelanin leads to lighter-skinned individuals having a pale pink skin tone. Individuals with lighter skin more readily absorb UV radiation as their basal melanin expression is directed more toward the production of pheomelanin than eumelanin. But why are there so many variations in skin tone in humans? To answer this question, we now turn toward an exploration of an evolutionary-based adaptation of skin tone as a function of the environment.<\/p>\n<h4 class=\"import-Normal\"><em>Adaptation: <\/em><em>Evolutionary Basis for Skin Tone Variation<\/em><\/h4>\n<figure style=\"width: 337px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image22-4.png\" alt=\"Table discussing hair and skin, folate and UV rays, leaving Africa, vitamin D, and selective pressures.\" width=\"337\" height=\"843\" \/><figcaption class=\"wp-caption-text\">Figure 15.12: Evolutionary basis for human skin colour variation. <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__-13\/\">Evolutionary basis for human skin color variation (Figure 14.12)<\/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>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Skin cancer is a significant concern for many individuals with light skin tone as the cumulative exposure of the epidermis and underlying skin tissues to UV radiation may result in the development of abnormal cells within those tissues, leading to malignancies. Although darker-skinned individuals are at risk for skin cancer as well, they are less likely to develop it due to increased levels of melanin, specifically eumelanin, in their skin. Even though skin cancer is a serious health concern for some individuals, most skin cancers occur in the postreproductive years; therefore, it is improbable that evolutionary forces favouring varying melanin expression levels are related to a selective pressure to avoid such cancers. Furthermore, if avoiding skin cancer were the primary factor driving the evolution of various skin tones, then it reasons that everyone would have the most significant expression of eumelanin possible. So, why do we have different skin tones (Figure 15.12)?<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The term <strong>cline<\/strong> (introduced in Chapter 14) refers to the continuum or spectrum of gradations (i.e., levels or degrees) from one extreme to another. With respect to skin tone, the various tonal shades occur clinally with darker skin being more prevalent near the equator and gradually decreasing in tone (i.e., decreased melanin production) in more distant latitudes. For individuals who are indigenous to equatorial regions, the increased levels of melanin within their skin provides them with a measure of protection against both sunburn and sunstroke because the melanin is more reflective of UV radiation than hemoglobin. In cases of severe sunburn, eccrine glands are affected, resulting in an individual\u2019s ability to sweat being compromised. As sweat is the body\u2019s most effective means of reducing its core temperature to maintain homeostasis, damage to the eccrine glands may lead to numerous physiological issues related to heat that may ultimately result in death.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Even though avoiding severe sunburn and sunstroke is of great importance to individuals within equatorial regions, this is likely not the primary factor driving the evolutionary selection of darker skin within these regions. It has been proposed that UV radiation\u2019s destruction of <strong>folic acid<\/strong>, which is a form of B-complex vitamin, may have led to the selection of darker skin in equatorial regions. For pregnant people, low levels of folic acid within the body during gestation may lead to defects in the formation of the brain and spinal cord of the fetus. This condition, which is referred to as spina bifida (Figure 15.13), often significantly reduces an infant\u2019s chances of survival without medical intervention. In people producing sperm, low levels of folic acid within the body reduce sperm quantity and quality. Thus, in geographic regions with high UV radiation levels (i.e., equatorial regions), there appears to be an evolutionarily driven correlation between darker skin and fertility.<\/p>\n<figure style=\"width: 364px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image19-2-1.jpg\" alt=\"Infant with a bulging spinal cord above the lower spine.\" width=\"364\" height=\"294\" \/><figcaption class=\"wp-caption-text\">Figure 15.13: Infant with open neural tube defect in lower (lumbar) region of the spine (right). A close-up of the open neural tube defect within the spinal column (left) shows the dura matter, which is ordinarily protected within the spine, is exposed on the surface of the skin. The spinal cord sits near the surface, when it should be protected within the spinal column. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Spina-bifida.jpg\">Spina-bifida<\/a> by the <a href=\"https:\/\/www.cdc.gov\/\">Centers for Disease Control and Prevention<\/a> is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p>If darker skin tone is potentially correlated to more successful reproduction, then why do lighter shades of skin exist? One hypothesis is that there is a relationship between lighter skin tone and vitamin D synthesis within the body. When skin is exposed to the UV-B radiation waves in sunlight, a series of chemical reactions occur within the epidermis leading to the production of vitamin D3, which is a fat-soluble vitamin that assists the body with absorbing calcium and phosphorus in the small intestine. These nutrients are among those that are critical for the proper growth and maintenance of bone tissue within the body. In the absence of adequate minerals, particularly calcium, bone structure and strength will be compromised, leading to the development of rickets during the growth phase. Rickets is a disease affecting children during their growth phase. It is characterized by inadequately calcified bones that are softer and more flexible than normal. Individuals with rickets will develop a true bowing of their legs, which may affect their mobility (Figure 15.14). In addition, deformation of pelvic bones in people who may become pregnant may occur as a result of rickets, leading to complications with reproduction. In adults, a deficiency in vitamin D3 will often result in osteomalacia, which is a general softening of the bones due to inadequate mineralization. As noted, a variety of maladies may occur due to the inadequate production or absorption of vitamin D3, as well as the destruction of folate within the human body. Therefore, from an evolutionary perspective, natural selection should favour a skin tone that is best suited to a given environment.<\/p>\n<figure style=\"width: 263px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image9-2-1.jpg\" alt=\"Historic photo of three young children each with visible lower limb curvature.\" width=\"263\" height=\"315\" \/><figcaption class=\"wp-caption-text\">Figure 15.14: Children with rickets in various developmental stages. Credit: <a href=\"https:\/\/wellcomecollection.org\/works\/m3eu5snb\">Rachitis, stages of development for children<\/a> [slide numbers 7181 and 7182; photo number: M0003399] by <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>In general, the trend related to lighter skin pigmentation further from the equator follows a principle called <strong>Gloger\u2019s Rule<\/strong>. This rule states that within the same species of mammals the more heavily pigmented individuals tend to originate near the equator while lighter-pigmented members of the species will be found in regions further from the equator. Gloger\u2019s Rule applies latitudinally; however, it does not appear to hold for certain human populations near the poles. Specifically, it does not apply to the Inuit people (Figure 15.15), who are indigenous to regions near the North Pole and currently reside in portions of Canada, Greenland, Alaska, and Denmark. The Inuit have a darker skin tone that would not be anticipated under the provisions of Gloger\u2019s Rule. The high reflectivity of light off of snow and ice, which is common in polar regions, necessitates the darker skin tone of these individuals to prevent folic acid degradation just as it does for individuals within equatorial regions. The consumption of vitamin D\u2013rich foods, such as raw fish, permits the Inuit to reside at high latitudes with darker skin tone while preventing rickets.<\/p>\n<h4 class=\"import-Normal\"><em>Adaptation: <\/em><em>Shape and Size Variations<\/em><\/h4>\n<figure style=\"width: 353px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image6-3-1.jpg\" alt=\"Historic photo of a person carrying a very large fish laid on a pack.\" width=\"353\" height=\"296\" \/><figcaption class=\"wp-caption-text\">Figure 15.15: Copper Inuk (from Bernard Harbour, Nunavut) with lake trout on his back (1915). Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Copper_Inuk_with_lake_trout_on_his_back_near_Bernard_Harbour_(51007).jpg\">Copper Inuk with lake trout on his back near Bernard Harbour, Northwest Territories (Nunavut)<\/a> (1915) by George H. Wilkins is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0 License<\/a>. This image is part of the Photographic Archives of the Canadian Museum of History.<\/figcaption><\/figure>\n<p>In addition to natural selection playing a role in the determination of melanin expression, it plays a significant role in the determination of the shape and size of the human body. As previously discussed, the most significant thermodynamic mechanism of heat loss from the body is radiation. At temperatures below 20\u2103 (68\u2109), the human body loses around 65% of its heat to radiative processes; however, the efficiency of radiation is correlated to the overall body shape and size of the individual. There is a direct correlation between the ratio of an object\u2019s surface area to mass and the amount of heat that may be lost through radiation. For example, two objects of identical composition and mass are heated to the same temperature. One object is a cube and the other is a sphere. Which object will cool the fastest? Geometrically, a sphere has the smallest surface area per unit mass of any three-dimensional object, so the sphere will cool more slowly than the cube. In other words, the smaller the ratio of the surface area to mass an object has, the more it will retain heat. With respect to the cube in our example, mass increases by the cube, but surface area may increase only by the square, so size will affect the mass to surface area ratio. This, in general, holds true for humans, as well.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In regions where temperatures are consistently cold, the body shape and size of individuals indigenous to the area tend to be more compact. These individuals have a relatively higher body mass to surface area (i.e., skin) than their counterparts from equatorial regions where the average temperatures are considerably warmer. Individuals from hot climates, such as the Fulani (Figure 15.16) of West Africa, have limbs that are considerably longer than those of individuals from cold climates, such as the Inuit of Greenland (Figure 15.16). Evolutionarily, the longer limbs of individuals from equatorial regions (e.g., the Fulani) provide a greater surface area (i.e., lower body mass to surface area ratio) for the dissipation of heat through radiative processes. In contrast, the relatively short limbs of Arctic-dwelling people, such as the Inuit, allows for the retention of heat because there is a decreased surface area through which heat may radiate away from the body.<\/p>\n<figure style=\"width: 385px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image20-2.jpg\" alt=\"Person on left is stocky, person on right has a narrow body breadth.\" width=\"385\" height=\"570\" \/><figcaption class=\"wp-caption-text\">Figure 15.16: The individual on the left is typical of one adapted to a cold environment where the conservation of heat in the body\u2019s core is of critical importance.The individual on the right could be adapted to a tropical environment where the rapid dispersal of heat is necessary to maintain homeostasis. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Greenland_1999_(01).jpg\">Greenland 1999 (01)<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Vadeve\">Vadeve<\/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\">As described above, there are certain trends related to the general shape and size of human bodies in relation to the thermal conditions. To better describe these trends, we turn to a couple of general principles that are applicable to a variety of species beyond humans. <strong>Bergmann\u2019s Rule<\/strong> predicts that as average environmental temperature decreases, populations are expected to exhibit an increase in weight and a decrease in surface area (Figure 15.17a). Also, within the same species of homeothermic animals, the relative length of projecting body parts (e.g., nose, ears, and limbs) increases in relation to the average environmental temperature (Figure 15.17b). This principle, referred to as <strong>Allen\u2019s Rule<\/strong>, notes that longer, thinner limbs are advantageous for the radiation of excess heat in hot environments and shorter, stockier limbs assist with the preservation of body heat in cold climates. A measure of the crural index (crural index = tibia length <span style=\"font-size: NaNpt;color: #;text-decoration: none\">\u00f7 <\/span>femur length) of individuals from various human populations provides support for Allen\u2019s Rule since this value is lower in individuals from colder climates than it is for those from hot climates. The crural indices for human populations vary directly with temperature, so individuals with higher crural index values are generally from regions with a warmer average environmental temperature. Conversely, the crural indices are lower for individuals from regions where there are colder average temperatures.<\/p>\n<figure style=\"width: 533px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image16-6.png\" alt=\"Illustration of an elk (left) and gazelle (right).\" width=\"533\" height=\"319\" \/><figcaption class=\"wp-caption-text\">Figure 15.17a: These organisms are representative of Bergmann\u2019s rule. The animal on the left depicts an ungulate from a cooler environment with increased body weight and decreased surface area, compared to the slender ungulate on the right. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-13\/\">Bergmann\u2019s Rule (Figure 14.16a)<\/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<figure style=\"width: 521px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image14-4.png\" alt=\"Three types of rabbits with different ear and forelimb lengths.\" width=\"521\" height=\"264\" \/><figcaption class=\"wp-caption-text\">Figure 15.17b: These animals are representative of Allen\u2019s rule. The rabbit on the left comes from a cooler environment and is compact with short limbs and ears. The rabbit on the right comes from a warm environment and has long and lanky limbs and ears. The rabbit in the middle has ears and limbs that are in-between the other two. Rabbits do not sweat like humans; heat is dissipated primarily through their ears. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-13\/\">Allen\u2019s Rule (Figure 14.16b)<\/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<figure style=\"width: 627px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image17-6.png\" alt=\"Four noses with varying heights, arches, and nostril breadth.\" width=\"627\" height=\"237\" \/><figcaption class=\"wp-caption-text\">Figure 15.18: Human nasal morphological variation as influenced by four major climate-based adaptive zones: hot-dry, hot-wet, cold-dry, and cold-wet. The four noses in this figure vary in shape in relation to their respective climate-based adaptive zones. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-13\/\">Human nasal morphological variation (Figure 14.17)<\/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>Nasal shape and size (Figure 15.18) is another physiological feature affected by our ancestors\u2019 environments. The selective role of climate in determining human nasal variation is typically approached by dividing climates into four adaptive zones: hot-dry, hot-wet, cold-dry, and cold-wet (Maddux Et al. 2016). A principal role of the nasal cavity is to condition (i.e., warm and humidify) ambient air prior to its reaching the lungs. Given this function of the nasal cavity, it is anticipated that different nasal shapes and sizes will be related to varying environments. In cold-dry climates, an individual\u2019s nasal cavity must provide humidification and warmth to the dry air when breathing in through the nose (Noback Et al. 2011). Also, in that type of climate, the nasal cavity must conserve moisture and minimize heat loss during when the individual exhales through the nose (Noback Et al. 2011). From a physiological stress perspective, this is a stressful event.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Conversely, in hot-wet environments, there is no need for the nasal cavity to provide additional moisture to the inhaled air nor is there a need to warm the air or to preserve heat within the nasal cavity (Noback Et al. 2011). So, in hot-wet climates, the body is under less physiological stress related to the inhalation of ambient air than in cold-dry climates. As with most human morphological elements, the shape and size of the nasal cavity occurs along a cline. Due to the environmental stressors of cold-dry environments requiring the humidification and warming of air through the nasal cavity, individuals indigenous to such environments tend to have taller (longer) noses with a reduced nasal entrance (nostril opening) size (Noback Et al. 2011). This general shape is referred to as leptorrhine, and it allows for a larger surface area within the nasal cavity itself for the air to be warmed and humidified prior to entering the lungs (Maddux Et al. 2016). In addition, the relatively small nasal entrance of leptorrhine noses serves as a means of conserving moisture and heat (Noback Et al. 2011). Individuals indigenous to hot-wet climates tend to have platyrrhine nasal shapes, which are shorter with broader nasal entrances (Maddux Et al. 2016). Since individuals in hot-wet climates do not need to humidify and warm the air entering the nose, their nasal tract is shorter and the nasal entrance wider to permit the effective cooling of the nasal cavity during respiratory processes.<\/p>\n<h4 class=\"import-Normal\"><em>Adaptation: Infectious Disease<\/em><\/h4>\n<p class=\"import-Normal\">Throughout our evolutionary journey, humans have been exposed to numerous infectious diseases. In the following section, we will explore some of the evolutionary-based adaptations that have occurred in certain populations in response to the stressors presented by select infectious diseases. One of the primary examples of natural selection processes acting on the human genome in response to the presence of an infectious disease is the case of the relationship between the sickle-cell anemia trait and malaria, introduced in Chapter 5.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Malaria is a zoonotic disease (an infectious disease transmitted between animals and humans; it is covered in more detail in Chapter 17). It is caused by the spread of the parasitic protozoa from the genus <em>Plasmodium <\/em>(Figure 15.19). These unicellular, eukaryotic protozoa are transmitted through the bite of a female <em>Anopheles<\/em> mosquito. During the bite process, the protozoan parasites present within an infected mosquito\u2019s saliva enter a host\u2019s bloodstream where they are transported to the liver. Within the liver, the parasites multiply and are eventually released into the bloodstream, where they infect erythrocytes. Once inside the erythrocytes, the parasites reproduce until they exceed the cell\u2019s storage capacity, causing it to burst and release the parasites into the bloodstream once again. This replication cycle continues as long as there are viable erythrocytes within the host to infect.<\/p>\n<figure style=\"width: 518px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image13-1-1.jpg\" alt=\"Life cycle stages of Malaria parasite.\" width=\"518\" height=\"511\" \/><figcaption class=\"wp-caption-text\">Figure 15.19: Life cycle of the malaria parasite. <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:Malaria_parasite_life_cycle-NIAID.jpg\">Malaria parasite life cycle-NIAID<\/a> by <a href=\"https:\/\/www.niaid.nih.gov\/\">NIH National Institute of Allergy and Infectious Diseases<\/a> is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">General complications from malaria infections include the following: enlargement of the spleen (due to destruction of infected erythrocytes); lower number of thrombocytes (also called platelets, required for coagulation\/clotting of blood); high levels of bilirubin (a byproduct of hemoglobin breakdown in the liver) in the blood; jaundice (yellowing of the skin and eyes due to increased blood bilirubin levels); fever; vomiting; retinal (eye) damage; and convulsions (seizures). In 2020, there were 241 million cases of malaria reported globally, with 95% of those cases originating in Africa (World Health Organization 2021). In sub-Saharan Africa, where incidents of malaria are the highest in the world, 125 million pregnancies are affected by malaria, resulting in 200,000 infant deaths (Hartman, Rogerson, and Fischer 2013). Pregnant people who become infected during the gestational process are more likely to have low-birthweight infants due to prematurity or growth restriction inside the uterus (Hartman, Rogerson, and Fischer 2013). After birth, infants born to malaria-infected pregnant people are more likely to develop infantile anemia (low red-blood cell counts), a malaria infection that is not related to the maternal malarial infection, and they are more likely to die than infants born to non-malaria-infected pregnant people (Hartman, Rogerson, and Fischer 2013).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">For children and adolescents whose brains are still developing, there is a risk of cognitive (intellectual) impairment associated with some forms of malaria infections (Fernando, Rodrigo, and Rajapakse 2010). Given the relatively high rates of morbidity (disease) and mortality (number of deaths) associated with malaria, it is plausible that this disease may have served as a selective pressure during human evolution. Support for natural selection related to malaria resistance is related to genetic mutations associated with sickle cell, thalassemia, glucose-6-phosphate dehydrogenase (G6PD) deficiency, and the absence of certain antigens (molecules capable of inducing an immune response from the host) on erythrocytes. For the purposes of this text, we will focus our discussion on the relationship between sickle cell disease and malaria.<\/p>\n<figure style=\"width: 312px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image15-2-1.jpg\" alt=\"Normal (round) and sickle (crescent-shaped) red blood cells flow through a blood vessel. \" width=\"312\" height=\"499\" \/><figcaption class=\"wp-caption-text\">Figure 15.20: Normal and sickled erythrocytes. Normal red blood cells are round and can easily flow freely through blood vessels. Abnormal, or cicle, blood cells are half moon shaped and can easily become entangled and block blood flow. Sickle cells have abnormal hemoglobin that form strands that cause the sickle shape. The Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Sickle_cell_01.jpg\">Sickle cell 01<\/a> by <a href=\"https:\/\/www.nhlbi.nih.gov\/health\/sickle-cell-disease\">The National Heart, Lung, and Blood Institute (NHLBI)<\/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\"><strong>Sickle cell disease<\/strong> is a group of genetically inherited blood disorders characterized by an abnormality in the shape of the hemoglobin within erythrocytes. It is important to note that there are multiple variants of hemoglobin, including, but not limited to the following: A, D, C, E, F, H, S, Barts, Portland, Hope, Pisa, and Hopkins. Each of these variants of hemoglobin may result in various conditions within the body; however, for the following explanation we will focus solely on variants A and S.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Individuals who inherit a mutated gene (hemoglobin with a sickled erythrocyte variety, HbS) on chromosome 11 from both parents will develop sickle cell anemia, which is the most severe form of the sickle cell disease family (Figure 15.20). The genotype of an individual with sickle cell anemia is HbSS; whereas, an individual without sickle cell alleles has a genotype of HbAA representing two normal adult hemoglobin type A variants. Manifestations of sickle cell anemia (HbSS) range from mild to severe, with some of the more common symptoms being anemia, blood clots, organ failure, chest pain, fever, and low blood-oxygen levels. In high-income countries with advanced medical care, the median life expectancy of an HbSS individual is around 60 years; however, in low-income countries where advanced medical care is scarce, as many as 90% of children with sickle cell disease perish before the age of five (Longo Et al. 2017).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Considering that advanced medical care was not available during much of human evolutionary history, it stands to reason that the majority of individuals with the HbSS genotype died before the age of reproduction. If that is the case though, why do we still have the HbS variant present in modern populations? As covered earlier in this textbook, the genotype of an individual is composed of genes from both biological parents. In the case of an individual with an HbSS genotype, the sickle cell allele (HbS) was inherited from each of the parents. For individuals with the heterozygous genotype of HbSA, they have inherited both a sickle cell allele (HbS) and a normal hemoglobin allele (HbA). Heterozygous (HbSA) individuals who reside in regions where malaria is endemic may have a selective advantage. They will experience a sickling of some, but not all, of their erythrocytes. As discussed in the following paragraph, HbSA heterozygous individuals are less likely to die from malaria infections than their HbAA counterparts. Unlike an individual with the HbSS genotype, someone with HbSA may experience some of the symptoms listed above; however, they are generally less severe.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">As noted earlier, the mechanism through which <em>Plasmodium<\/em> protozoan parasites replicate involves human erythrocyte cells. However, due to their sickled shape, as well as the presence of an abnormally shaped protein within the cell, the parasites are unable to replicate effectively in the erythrocyte cells coded for by the HbS allele (Cyrklaff Et al. 2011). An individual who has an HbSA genotype and an active malaria infection will become ill with the disease to a lesser extent than someone with an HbAA genotype, which increases their chances of survival. Although normal erythrocytes (regulated by the HbA allele) allow for parasite replication, they are not able to replicate in HbS erythrocytes of the heterozygote. So, individuals with the HbSA genotype are more likely to survive a malaria infection than an individual who is HbAA. Although individuals with the HbSA genotype may endure some physiological complications related to the sickling of some of their erythrocytes, their morbidity and mortality rates are lower than they are for HbSS members of the population. The majority of individuals who are heterozygous or homozygous for the HbS trait have ancestors who originated in sub-Saharan Africa, India, Saudi Arabia, and regions in South and Central America, the Mediterranean (Turkey, Greece, and Italy), and the Caribbean (Centers for Disease Control and Prevention 2017; Figure 15.21).<\/p>\n<figure style=\"width: 511px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image23-3.png\" alt=\"Map illustrating distribution of sickle cell and associated erythrocytic abnormalities for Africa and Asia.\" width=\"511\" height=\"306\" \/><figcaption class=\"wp-caption-text\">Figure 15.21: Distribution of sickle cell and associated erythrocytic abnormalities for Africa and Asia. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Red_Blood_Cell_abnormalities.png\">Red Blood Cell abnormalities<\/a> by Armando Moreno Vranich 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\">With respect to the history of these regions, during the early phases of settlement horticulture was the primary method of crop cultivation. Typically performed on a small scale, horticulture is based on manual labour and relatively simple hand tools rather than the use of draft animals or irrigation technologies. Common in horticulture is <em>swidden<\/em>, or the cutting and burning of plants in woodland and grassland regions. The swidden is the prepared field that results following a slash-and-burn episode. This practice fundamentally alters the soil chemistry, removes plants that provide shade, and increases the areas where water may pool. This anthropogenically altered landscape provides the perfect breeding ground for the <em>Anopheles<\/em> mosquito, as it prefers warm, stagnant pools of water (Figure 15.22).<\/p>\n<figure style=\"width: 561px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5-6.png\" alt=\"Horticulture encourages mosquitos, increasing malaria. Balancing selection maintains normal and sickle cell alleles.\" width=\"561\" height=\"468\" \/><figcaption class=\"wp-caption-text\">Figure 15.22: The effects of human horticultural activities on the balancing selection of populations in relation to sickle cell disease genotype variants. <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__-13\/\">Sickle cell disease (Figure 14.21)<\/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>. [Includes two horticulture illustrations by Mary Nelson, <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Sickle_cell_anemia.jpg\">Sickle cell anemia<\/a> by <a href=\"https:\/\/en.wikibooks.org\/wiki\/User:Pkleong\">Pkleong<\/a> at <a href=\"https:\/\/en.wikibooks.org\/wiki\/\">English Wikibooks<\/a> (modified), <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\">Although swidden agriculture was historically practiced across the globe, it became most problematic in the regions where the <em>Anopheles<\/em> mosquito is endemic. These areas have the highest incidence rates of malaria infection. Over time, the presence of the <em>Anopheles<\/em> mosquito and the <em>Plasmodium<\/em> parasite that it transmitted acted as a selective pressure, particularly in regions where swidden agricultural practices were common, toward the selection of individuals with some modicum of resistance against the infection. In these regions, HbSS and HbSA individuals would have been more likely to survive and reproduce successfully. Although individuals and populations are far more mobile now than they have been throughout much of history, there are still regions where we can see higher rates of malaria infection as well as greater numbers of individuals with the HbS erythrocyte variant. The relationship between malaria and the selective pressure for the HbS variant is one of the most prominent examples of natural selection in the human species within recent evolutionary history.<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><em>Adaptation: Lactase Persistence <\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">With the case of sickled erythrocytes and their resistance to infection by malaria parasites, there is strong support for a cause-and-effect-style relationship linked to natural selection. Although somewhat less apparent, there is a correlation between lactase persistence and environmental challenges. Lactase-phlorizin hydrolase (LPH) is an enzyme that is primarily produced in the small intestine and permits the proper digestion of lactose, a disaccharide (composed of two simple sugars: glucose and galactose) found in the milk of mammals. Most humans will experience a decrease in the expression of LPH following weaning, leading to an inability to properly digest lactose. Generally, LPH production decreases between the ages of two and five and is completely absent by the age of nine (Dzialanski Et al. 2016). For these individuals, the ingestion of lactose may lead to a wide variety of gastrointestinal ailments, including abdominal bloating, increased gas, and diarrhea. Although the bloating and gas are unpleasant, the diarrhea caused by a failure to properly digest lactose can be life-threatening if severe enough due to the dehydration it can cause. Some humans, however, are able to produce LPH far beyond the weaning period.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Individuals who continue to produce LPH have what is referred to as the <strong>lactase persistence <\/strong>trait. The lactase persistence trait is encoded for a gene called <em>LCT<\/em>, which is located on human chromosome 2 (Ranciaro Et al. 2014). From an evolutionary and historical perspective, this trait is most commonly linked to cultures that have practiced cattle domestication (Figure 15.23). For individuals in those cultures, the continued expression of LPH may have provided a selective advantage. During periods of environmental stress, such as a drought, if an individual is capable of successfully digesting cow\u2019s milk, they have a higher chance of survival than someone who suffers from diarrhea-linked dehydration due to a lack of LPH. Although the frequency of the lactase persistence trait is relatively low among African agriculturalists, it is high among pastoralist populations that are traditionally associated with cattle domestication, such as the Tutsi and Fulani, who have frequencies of 90% and 50%, respectively (Tishkoff Et al. 2007).<\/p>\n<figure style=\"width: 632px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-4.png\" alt=\"Map depicting the percentage of adults with lactase persistence in the eastern hemisphere.\" width=\"632\" height=\"497\" \/><figcaption class=\"wp-caption-text\">Figure 15.23: Interpolated map depicting the percentage of adults with the lactase persistence genotype in indigenous populations of Africa, Europe, Asia, and Australia. Circles denote sample locations. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Lactose_tolerance_in_the_Old_World.svg\">Lactose tolerance in the Old World<\/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\/4.0\/legalcode\">CC BY 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Cattle domestication began around 11,000 years ago in Europe (Beja-Pereira Et al. 2006) and 7,500 to 9,000 years ago in the Middle East and North Africa (Tishkoff Et al. 2007). Based on human genomic studies, it is estimated that the mutation for the lactase persistence trait occurred around 2,000 to 20,000 years ago for European populations (Tishkoff Et al. 2007). For African populations, the lactase persistence trait emerged approximately 1,200 to 23,000 years ago (Gerbault Et al. 2011). This begs the question: Is this mutation the same for both populations? It appears that the emergence of the lactase persistence mutation in non-European populations, specifically those in East Africa (e.g., Tutsi and Fulani), is a case of <strong>convergent evolution<\/strong>. With convergent evolution events, a similar mutation may occur in species of different lineages through independent evolutionary processes. Based on our current understanding of the genetic mutation pathways for the lactase persistence trait in European and African populations, these mutations are not representative of a shared lineage. In other words, just because a person of European origin and a person of African origin can each digest milk due to the presence of the lactase-persistence trait in their genotypes, it does not mean that these two individuals inherited it due to shared common ancestry.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Is it possible that the convergent evolution of similar lactase-persistence traits in disparate populations is merely a product of genetic drift? Or is there evidence for natural selection? Even though 23,000 years may seem like a long time, it is but a blink of the proverbial evolutionary eye. From the perspective of human evolutionary pathways, mutations related to the <em>LCT<\/em> gene have occurred relatively recently. Similar genetic changes in multiple populations through genetic drift processes, which are relatively slow and directionless, fail to accumulate as rapidly as lactase-persistence traits (Gerbault Et al. 2011). The widespread accumulation of these traits in a relatively short period of time supports the notion that an underlying selective pressure must be driving this form of human evolution. Although to date no definitive factors have been firmly identified, it is thought that environmental pressures are likely to credit for the rapid accumulation of the lactase-persistence trait in multiple human populations through convergent evolutionary pathways.<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Skin Tone Genetic Regulation<\/h2>\n<p class=\"import-Normal\">The melanocortin 1 receptor (<em>MC1R<\/em>) gene acts to control which types of melanin (eumelanin or pheomelanin) are produced by melanocytes. The <em>MC1R<\/em> receptor is located on the surface of the melanocyte cells (Quillen Et al. 2018). Activation of the <em>MC1R <\/em>receptors may occur through exposure to specific environmental stimuli or due to underlying genetic processes. Inactive or blocked <em>MC1R <\/em>receptors result in melanocytes producing pheomelanin. If the <em>MC1R<\/em> gene receptors are activated, then the melanocytes will produce eumelanin. Thus, individuals with activated <em>MC1R<\/em> receptors tend to have darker-pigmented skin and hair than individuals with inactive or blocked receptors.<\/p>\n<p class=\"import-Normal\">The alleles of another gene, the major facilitator, superfamily domain-containing protein 12 (<em>MFSD12<\/em>) gene, affect the expression of melanocytes in a different way than the <em>MC1R<\/em> gene. Instead of affecting the activation of melanocyte receptors, the <em>MFSD12<\/em> alleles indirectly affect the membranes of melanocyte lysosomes (Quillen Et al. 2018). The melanocyte\u2019s lysosomes are organelles containing digestive enzymes, which ultimately correlate to varying degrees of pigmentation in humans. Variations in the membranes of the melanocyte lysosomes ultimately correlate to differing degrees of pigmentation in humans.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ancestral <em>MFSD12 <\/em>allele variants are present in European and East Asian populations and are associated with lighter pigmentation of the skin (Crawford Et al. 2017; Quillen Et al. 2018). In addition, this ancestral variant is also associated with Tanzanian, San, and Ethiopian populations of Afro-Asiatic ancestry (Crawford Et al. 2017; Quillen Et al. 2018). In contrast, the more derived (i.e., more recent) allele variants that are linked to darker skin tones are more commonly present in East African populations, particularly those of Nilo-Saharan descent (Crawford Et al. 2017; Quillen Et al. 2018). The notion that ancestral alleles of <em>MFSD12<\/em> are associated with lighter skin pigmentation is in opposition to the commonly accepted idea that our pigmentation was likely darker throughout early human evolution (Crawford Et al. 2017; Quillen Et al. 2018). Due to the complexity of the human genome, <em>MFSD12<\/em> and <em>MC1R <\/em>are but two examples of alleles affecting human skin tone. Furthermore, there is genetic evidence suggesting that certain genomic variants associated with both darker and lighter skin colour have been subject to directional selection processes for as long as 600,000 years, which far exceeds the evolutionary span of <em>Homo sapiens sapiens<\/em> (Crawford Et al. 2017; Quillen Et al. 2018).<\/p>\n<\/div>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<div class=\"textbox shaded\">\n<h2>Summary<\/h2>\n<p>From the time that the first of our species left Africa, we have had to adjust and adapt to numerous environmental challenges. The remarkable ability of human beings to maintain homeostasis through a combination of both nongenetic (adjustments) and genetic (adaptations) means has allowed us to occupy a remarkable variety of environments, from high-altitude mountainous regions to the tropics near the equator. From adding piquant, pungent spices to our foods as a means of inhibiting food-borne illnesses due to bacterial growth to donning garments specially suited to local climates, behavioural adjustments have provided us with a nongenetic means of coping with obstacles to our health and well-being. Acclimatory adjustments, such as sweating when we are warm in an attempt to regulate our body temperature or experiencing increased breathing rates as a means of increasing blood oxygen levels in regions where the partial pressure of oxygen is low, have been instrumental in our survival with respect to thermal and altitudinal environmental challenges. For some individuals, developmental adjustments that were acquired during their development and growth phases (e.g., increased heart and lung capacities for individuals from high-altitude regions) provide them with a form of physiological advantage not possible for someone who ventures to such an environmentally challenging region as an adult. Genetically mediated adaptations, such as variations in the pigmentation of our skin, have ensured our evolutionary fitness across all latitudes. Taken together, these examples point to a broader question: will the human species continue to adjust and adapt to new environmental challenges in the future? If past performance is any measure of future expectations, then the human story will continue as long as we do not alter our environment to the point that the plasticity of our behaviour, physiological, and morphological boundaries is exceeded.<\/p>\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\">Detail at least two examples of how natural selection has influenced human variation. Specifically, what was the selective pressure that may have led to a preference for a specific trait and how is that trait related to an increased level of fitness?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Why is reduced pigmentation of the skin advantageous for individuals from northern latitudes? What role does darker skin pigmentation serve for individuals near the equator? What is the relationship between skin pigmentation and fitness?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">What are some of the risks associated with pregnancy at high altitude? Compare and contrast the various genetic mutations of the indigenous Tibetan and Ethiopian high-altitude populations. In your answer, specifically address the issue of pregnancy at high altitudes.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">What is the relationship between the sickle cell mutation and the <em>Plasmodium<\/em> parasite? Would having the HbSA genotype still be advantageous in a region where such parasites are not common? Why or why not?<\/li>\n<\/ul>\n<\/div>\n<p>&nbsp;<\/p>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Key Terms<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Acclimatory adjustm<\/strong><strong>ents<\/strong>: Processes by which an individual organism adjusts in order to maintain homeostasis in response to environmental challenges.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Activated melanogenesis<\/strong>: Increase in melanin production in response to ultraviolet radiation (UV) exposure.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Adaptation<\/strong>: Alteration in population-level gene frequencies related to environmentally induced selective pressures; leads to a greater level of fitness for a population related to a specific environment.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Adjustment<\/strong>: Nongenetic-based ways in which organisms adjust to environmental stressors.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Allen\u2019s Rule<\/strong>: Due to thermal adaptation, homeothermic animals have body volume-to-surface ratios that vary inversely with the average temperature of their environment. In cold climates, the anticipated ratio is high; in warm climates, it is low.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Basal melanogenesis<\/strong>: Genetically mediated, non-environmentally influenced base melanin level.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Behavioural adjustments<\/strong>: An individual\u2019s culturally mediated responses to an environmental stressor in an effort to maintain homeostasis.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Bergmann\u2019s Rule<\/strong>: For a broadly distributed monophyletic group, species and populations of smaller size tend to be found in environments with warmer climates and those of larger size tend to be found in ones that are colder.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Cline<\/strong>: A continuum of gradations (i.e., degrees or levels) of a specific trait.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Conduction<\/strong>: Mechanism of heat transfer between objects through direct contact.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Convection<\/strong>: Movement of heat away from a warm object to the cooler surrounding fluid (i.e., gas or liquid).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Convergent evolution<\/strong>: Evolutionary process whereby organisms that are not closely related independently evolve similar traits as a product of adaptation to similar evolutionary parameters.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Erythrocyte<\/strong>: Red blood cell; most common form of blood cell; the principle means of transporting oxygen throughout the circulatory system.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Evaporation<\/strong>: Mechanism of heat transfer whereby liquid is transformed into a gas, utilizing energy (e.g., heat).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Folic acid<\/strong>: Form of B complex vitamin necessary for proper fetal development.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Gloger\u2019s Rule<\/strong>: For mammals of the same species, those with more darkly pigmented forms tend to be found closer to the equator and those with lighter forms are found in regions further from the equator.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Homeostasis<\/strong>: Condition of optimal functioning for an organism.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Hyperpnea<\/strong>: Increased depth and rate of respiration.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Hypothalamus<\/strong>: Small portion of the human brain responsible for body temperature regulation.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Lactase persistence<\/strong>: Genetic mutation permitting the continued production of lactase-phlorizin hydrolase enzyme in the small intestine past the weaning period.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Melanin<\/strong>: Black-brown pigment produced by melanocytes; one of the primary pigments in skin.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Melanocytes<\/strong>: Specialized cells that produce melanin.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Phenotypic plasticity<\/strong>: Ability of one genotype to produce more than one phenotype dependent on environmental conditions.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Radiation<\/strong>: Mechanism of heat transfer involving electromagnetic energy being emitted from an object.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Sickle cell disease<\/strong>: A group of genetically inherited blood disorders characterized by an abnormality in the shape of the hemoglobin within erythrocytes (red blood cells).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Stressor<\/strong>: Any stimulus resulting in an imbalance in an organism\u2019s homeostatic balance.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Vasoconstriction<\/strong>: Narrowing of the blood vessels due to contractions of the muscular vessel walls.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Vasodilation<\/strong>: Dilation of the blood vessels due to relaxation of the muscular vessel walls.<\/p>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">For Further Exploration<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Homeostasis<\/strong><\/p>\n<p class=\"import-Normal\">Baptista, Vander. 2006. \u201c<a href=\"https:\/\/doi.org\/10.1152\/advan.00075.2006\" target=\"_blank\" rel=\"noopener\">Starting Physiology: Understanding Homeostasis<\/a>.\u201d <em>Advances in Physiology Education<\/em> 30: 263\u2013264.<\/p>\n<p class=\"import-Normal\">Goldstein, David S., and Bruce McEwen. 2002. \u201c<a href=\"https:\/\/doi.org\/10.1080\/102538902900012345\" target=\"_blank\" rel=\"noopener\">Allostasis, Homeostats, and the Nature of Stress<\/a>.\u201d <em>The International Journal on the Biology of Stress<\/em> 5 (1): 55\u201358.<\/p>\n<p class=\"import-Normal\"><strong>General Clinal Variation and Genetic Exchange<\/strong><\/p>\n<p class=\"import-Normal\">Delhey, Kaspar. 2019. \u201c<a href=\"https:\/\/doi.org\/10.1111\/brv.12503\" target=\"_blank\" rel=\"noopener\">A Review of Gloger's Rule, an Ecogeographical Rule of Colour: Definitions, Interpretations and Evidence<\/a>.\u201d <em>Biological Reviews<\/em> 94 (4): 1294\u20131316.<\/p>\n<p class=\"import-Normal\">Feng, Yuanqing, Michael A. McQuillan, and Sarah A. Tishkoff. 2021. \u201c<a href=\"https:\/\/doi.org\/10.1093\/hmg\/ddab007\" target=\"_blank\" rel=\"noopener\">Evolutionary Genetics of Skin Pigmentation in African Populations<\/a>.\u201d <em>Human Molecular Genetics<\/em> 30 (R1): R88\u2013R97.<\/p>\n<p class=\"import-Normal\">Hu, Hao, Nayia Petousi, Gustavo Glusman, Yao Yu, Ryan Bohlender, Tsewang Tashi, Jonathan M. Downie, et al. 2017. \u201c<a href=\"https:\/\/doi.org\/10.1371\/journal.pgen.1006675\" target=\"_blank\" rel=\"noopener\">Evolutionary History of Tibetans Inferred from Whole-Genome Sequencing<\/a>.\u201d <em>PLoS Genetics <\/em>13 (4): e1006675. .<\/p>\n<p class=\"import-Normal\">Jablonski, Nina G. 2021. \u201c<a href=\"https:\/\/doi.org\/10.1002\/ajpa.24200\" target=\"_blank\" rel=\"noopener\">Skin Color and Race<\/a>.\u201d Special issue, \u201cRace Reconciled II: Interpreting and Communicating Biological Variation and Race in 2021,\u201d <em>American Journal of Physical Anthropology<\/em> 175 (2): 437\u2013447.<\/p>\n<p class=\"import-Normal\">Pritchard, Jonathan K., Joseph K. Pickrell, and Graham Coop. 2010. \u201c<a href=\"https:\/\/doi.org\/10.1016\/j.cub.2009.11.055\" target=\"_blank\" rel=\"noopener\">The Genetics of Human Adaptation: Hard Sweeps, Soft Sweeps, and Polygenic Adaptation<\/a>.\u201d <em>Current Biology<\/em> 20 (4): R208\u2013R215.<\/p>\n<p class=\"import-Normal\">Sankararaman, Sriram, Swapan Mallick, Nick Patterson, and David Reich. 2016. \u201c<a href=\"https:\/\/doi.org\/10.1016\/j.cub.2016.03.037\" target=\"_blank\" rel=\"noopener\">The Combined Landscape of Denisovan and Neanderthal Ancestry in Present-Day Humans<\/a>.\u201d <em>Current Biology<\/em> 26 (9): 1241\u20131247.<\/p>\n<p class=\"import-Normal\"><strong>Lactase Persistence<\/strong><\/p>\n<p class=\"import-Normal\">HHMI BioInteractive. 2021. \u201c<a href=\"https:\/\/www.biointeractive.org\/classroom-resources\/making-fittest-got-lactase-co-evolution-genes-and-culture\" target=\"_blank\" rel=\"noopener\">The Making of the Fittest: Got Lactase? The Co-evolution of Genes and Culture<\/a>.\u201d Accessed April 7, 2023.<\/p>\n<p class=\"import-Normal\"><strong>Malaria and Sickle Cell Anemia<\/strong><\/p>\n<p class=\"import-Normal\">Bill and Melinda Gates Foundation. 2022. \u201c<a href=\"https:\/\/www.gatesfoundation.org\/our-work\/programs\/global-health\/malaria\" target=\"_blank\" rel=\"noopener\">Malaria<\/a>.\u201d Accessed April 7, 2023.<\/p>\n<p class=\"import-Normal\">Centers for Disease Control and Prevention. 2022. \u201c<a href=\"https:\/\/www.cdc.gov\/parasites\/malaria\/index.html\" target=\"_blank\" rel=\"noopener\">Malaria<\/a>.\u201d Accessed April 7, 2023.<\/p>\n<p class=\"import-Normal\">HHMI BioInteractive. 2020. \u201c<a href=\"https:\/\/www.biointeractive.org\/classroom-resources\/making-fittest-natural-selection-humans\" target=\"_blank\" rel=\"noopener\">The Making of the Fittest: Natural Selection in Humans<\/a>.\u201d 2020. Accessed April 7, 2023.<\/p>\n<p class=\"import-Normal\">National Institutes of Health: National Center for Advancing Translational Sciences. \u201c<a href=\"https:\/\/rarediseases.info.nih.gov\/diseases\/8614\/sickle-cell-anemia\" target=\"_blank\" rel=\"noopener\">Sickle Cell Anemia<\/a>.\u201d Accessed April 7, 2023.<\/p>\n<p class=\"import-Normal\">World Health Organization. 2022. \u201c<a href=\"https:\/\/www.who.int\/news-room\/fact-sheets\/detail\/malaria\" target=\"_blank\" rel=\"noopener\">Malaria<\/a>.\u201d Accessed April 7, 2023.<\/p>\n<p class=\"import-Normal\"><strong>Rickets and Bone Health<\/strong><\/p>\n<p class=\"import-Normal\">National Institutes of Health: National Center for Advancing Translational Sciences. \u201c<a href=\"https:\/\/rarediseases.info.nih.gov\/diseases\/5700\/rickets\" target=\"_blank\" rel=\"noopener\">Rickets<\/a>.\u201d Accessed April 7, 2023.<\/p>\n<p class=\"import-Normal\">Talmadge, D. W., and R. V. Talmadge. 2007. \u201cCalcium Homeostasis: How Bone Solubility Relates to All Aspects of Bone Physiology.\u201d <em>Journal of Musculoskeletal and Neuronal Interactions <\/em>7 (2): 108\u2013112.<\/p>\n<p class=\"import-Normal\"><strong>Skin Colour<\/strong><\/p>\n<p class=\"import-Normal\">HHMI BioInteractive. 2020. \u201c<a href=\"https:\/\/www.biointeractive.org\/classroom-resources\/biology-skin-color\" target=\"_blank\" rel=\"noopener\">The Biology of Skin Color<\/a>.\u201d Accessed April 7, 2023<\/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\">American Academy of Pediatrics, Task Force on Infant Sleep Position and Sudden Infant Death Syndrome. 2000. \u201cChanging Concepts of Sudden Infant Death Syndrome: Implications for Infant Sleeping Environment and Sleep Position.\u201d <em>Pediatrics<\/em> 105 (3): 650\u2013656.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Beja-Pereira, Albano, David Caramelli, Carles Lalueza-Fox, Cristiano Vernesi, Nuno Ferrand, Antonella Casoli, Felix Goyache, et al. 2006. \u201cThe Origin of European Cattle: Evidence from Modern and Ancient DNA.\u201d <em>PNAS <\/em>103 (21): 8113\u20138118.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Best, Andre, Daniel E. Lieberman, and Jason M. Kamilar. 2019. \u201cDiversity and Evolution of Human Eccrine Sweat Gland Density.\u201d <em>Journal of Thermal Biology<\/em> 84: 331\u2013338.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Bigham, Abigail W. 2016. \u201c<a href=\"https:\/\/www.cdc.gov\/ncbddd\/sicklecell\/data.html\" target=\"_blank\" rel=\"noopener\">Genetics of Human Origin and Evolution: High-Altitude Adaptations<\/a>.\u201d <em>Current Opinion in Genetics &amp; Development<\/em> 41: 8\u201313.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Centers for Disease Control and Prevention. 2017. \u201cData &amp; Statistics on Sickle Cell Disease.\u201d <em>Centers for Disease Control and Prevention<\/em> website, August 9. Accessed April 7, 2023. .<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Crawford, Nicholas G., Derek E. Kelly, Matthew E. B. Hansen, Marcia H. Beltrame, Shaohua Fan, Shanna L. Bowman, Ethan Jewett, et al. 2017. \u201cLoci Associated with Skin Pigmentation Identified in African Populations.\u201d <em>Science<\/em> 358 (6365): 1\u201349.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Cyrkalff, Marek, Cecilia P. Sanchez, Nicole Kilian, Curille Bisseye, Jacques Simpore, Friedrich Frischknecht, and Michael Lanzer. 2011. \u201cHemoglobins S and C Interfere with Actin Remodeling in <em>Plasmodium <\/em><em>falciparum<\/em>-Infected Erythrocytes.\u201d <em>Science<\/em> 334 (6060): 1283\u20131286.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Dzialanski, Zbigniew, Michael Barany, Peter Engfeldt, Anders Magnuson, Lovisa A. Olsson, and Torbj\u04e7rn K. Nilsson. 2016. \u201cLactase Persistence versus Lactose Intolerance: Is There an Intermediate Phenotype?\u201d <em>Clinical Biochemistry<\/em> 49 (2016): 248\u2013252.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Fernando, Sumadya D., Chaturaka Rodrigo, and Senaka Rajapakse. 2010. \u201cThe \u2018Hidden\u2019 Burden of Malaria: Cognitive Impairment Following Infection.\u201d <em>Malaria Journal<\/em> 9 (366): 1\u201311.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Gerbault, Pascale, Anke Liebert, Yuval Itan, Adam Powell, Mathias Currat, Joachim Burger, Dallas M. Swallow, and Mark G. Thomas. 2011. \u201cEvolution of Lactase Persistence: An Example of Human Niche Construction.\u201d <em>Philosophical Transactions of the Royal Society B: Biological Sciences <\/em>366 (1566): 863\u2013877.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Gray, Olivia A., Jennifer Yoo, D\u00e9bora R. Sobriera, Jordan Jousma, David Witnosky, Noboru J. Sakabe, Ying-Jie Ping, et al. 2022. \u201cA Pleiotropic Hypoxia-Sensitive <em>EPAS1<\/em> Enhancer Is Disrupted by Adaptive Alleles in Tibetans.\u201d <em>Science Advances <\/em>8 (47): 1\u201313.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Hartman, T. K., S. J. Rogerson, and P. R. Fischer. 2013. \u201cThe Impact of Maternal Malaria on Newborns.\u201d <em>Annals of Tropical Paediatrics<\/em> 30 (4): 271\u2013282.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Longo, Dan L., Fr\u00e9d\u00e9ric B. Piel, Martin H. Steinberg, and David C. Rees. 2017. \u201cSickle Cell Disease.\u201d <em>The New England Journal of Medicine<\/em> 376 (16): 1561\u20131573.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Maddux, Scott D., Todd R. Yokley, Bohumil M. Svoma, and Robert G. Franciscus. 2016. \u201cAbsolute Humidity and the Human Nose: A Reanalysis of Climate Zones and Their Influence on Nasal Form and Function.\u201d <em>American Journal of Physical Anthropology <\/em>161 (2): 309\u2013320.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Meyer, M. C., M. S. Alexander, Z. Wang, D. L. Hoffmann, J. A. Dahl, D. Degering, W. R. Haas, and F. Schl\u00fctz. 2017. \u201cPermanent Human Occupation of the Central Tibetan Plateau in the Early Holocene.\u201d <em>Science <\/em>355 (6320): 64\u201367.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Moore, Lorna G., Susan Niermeyer, and Stacy Zamudio. 1998. \u201cHuman Adaptation to High Altitude: Regional and Life-Cycle Perspectives.\u201d <em>Yearbook of Physical Anthropology<\/em> 41: 25\u201364.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Noback, Marlijn L., Katerina Harvati, and Fred Spoor. 2011. \u201cClimate-Related Variation of the Human Nasal Cavity.\u201d <em>American Journal of Physical Anthropology<\/em> 145 (4): 599\u2013614.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Peacock, A. J. 1998. \u201cABC of Oxygen: Oxygen at High Altitude.\u201d <em>BMJ<\/em> 317 (7165): 1063\u20131066.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Pontzer, Herman, Mary H. Brown, Brian M. Wood, David A. Raichlen, Audax Z.P. Madbulla, Jacob A. Harris, Holly Dunsworth, et al. 2021. \u201cEvolution of Water Conservation in Humans.\u201d <em>Current Biology<\/em> 31 (8): 1804\u20131810.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Quillen, Ellen E., Heather L. Norton, Esteban J. Parra, Frida Loza-Durazo, Khai C. Ang, Florin Mircea Illiescu, Laurel N. Pearson, et al. 2019. \u201cShades of Complexity: New Perspectives on the Evolution and Genetic Architecture of Human Skin.\u201d <em>Yearbook<\/em><em> of Physical Anthropology <\/em>168 (S67): 4\u201326.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ranciaro, Alessia, Michael C. Campbell, Jibril B. Hirbo, Wen-Ya Ko, Alain Froment,<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Paolo Anagnostou, Maritha J. Kotze, et al. 2014. \u201cGenetic Origins of Lactase Persistence and the Spread of Pastoralism in Africa.\u201d <em>American Journal of Human Genetics <\/em>94 (4): 496\u2013510.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Roby, Brianne Barnett, Marsha Finkelstein, Robert J. Tibesar, and James D. Sidman. 2012. \u201cPrevalence of Positional Plagiocephaly in Teens Born after the \u2018Back to Sleep\u2019 Campaign.\u201d <em>Otolaryn<\/em><em>g<\/em><em>ology<\/em><em>\u2014Head and Neck Surgery<\/em> 146 (5): 823\u2013828.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Sherman, Paul W., and Jennifer Billing. 1999. \u201cDarwinian Gastronomy: Why We Use Spices.\u201d <em>BioScience <\/em>49 (6): 453\u2013463.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Tishkoff, Sarah A., Floyd A. Reed, Alessia Ranciaro, Benjamin F. Voight, Courtney C. Babbitt, Jesse S. Silverman, Kweli Powell, et al. 2007. \u201cConvergent Adaptation of Human Lactase Persistence in Africa and Europe.\u201d <em>Nature Genetics<\/em> 39 (1): 31\u201340.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">World Health Organization. 2021. \u201c<a href=\"https:\/\/www.who.int\/teams\/global-malaria-programme\/reports\/world-malaria-report-2021\" target=\"_blank\" rel=\"noopener\">World Malaria Report 2021<\/a>.\u201d <em>World Health Organization<\/em> website, December 4, 2022. Accessed April 7, 2023.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Zhang, Xinjun, Kelsey E. Witt, Mayra M. Ba\u00f1uelos, Amy Ko, Kai Yuan, Shuhua Xu, Rasmus Nielsen, and Emilia Huerta-Sanchez. 2021. \u201cThe History and Evolution of Denisovan-<em>EPAS1<\/em> Haplotype in Tibetans.\u201d <em>PNAS Biological Sciences<\/em> 118 (22): 1\u20139.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">\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_1683_1723\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1723\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<div class=\"textbox\">\n<p>Learn more about <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/about-the-authors\/\">authors and editors<\/a>,<a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/student-contributors\/\"> Hess' student contributions<\/a>, and <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/versioning-history\/\">versioning history<\/a>.<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\">Learning Objectives<\/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 and 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-coloured 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 11.1). 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><img class=\"aligncenter\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image13-1.jpg\" alt=\"Reconstructed head and shoulders of a young Homo erectus.\" width=\"405\" height=\"308\" \/><figcaption class=\"wp-caption-text\">Figure 11.1a-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 10 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<\/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<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 8, 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 10, 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_1683_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_1683_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 11.2, 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 11.2: 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. It is speculated that 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 11.3) 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<figure style=\"width: 583px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 11.3: 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 undoubtedly challenging for our ancestors, but it offered new opportunities to make a living. It is theorized that 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 10) likely developed their large molar teeth with thick enamel in order to exploit this particular dietary niche.<\/span><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><span style=\"color: #000000\">Members of the genus <em>Homo <\/em>took a different route. Faced with the unstable African climate and shifting landscape, it is proposed that they evolved bigger brains that might have enabled them to rely on cultural solutions such as crafting stone tools that opened up new foraging opportunities. This strategy of behavioural flexibility might have served them well during this unpredictable time and may have 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 centimetres (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<figure style=\"width: 554px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image16.jpg\" alt=\"Africa map with South Africa, Tanzania, Kenya, and Ethiopia shaded.\" width=\"554\" height=\"717\" \/><figcaption class=\"wp-caption-text\">Figure 11.4: 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<p class=\"import-Normal\"><span style=\"color: #000000\">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 11.4). 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 analyse the remains. The fossil of the juvenile shown in Figure 11.5 (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<\/div>\n<figure id=\"attachment_316-2\" aria-describedby=\"caption-attachment-316-2\" style=\"width: 641px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-316\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/10.5.jpg\" alt=\"Two lateral right views of skulls and a jaw with blackened teeth.\" width=\"641\" height=\"214\" \/><figcaption id=\"caption-attachment-316-2\" class=\"wp-caption-text\">Figure 11.5a-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 the 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 11.6). 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 11.7). Researchers who favoured keeping all fossils in <em>Homo habilis<\/em> argued that sexual dimorphism, adaptation to local environments, or<strong> <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1566\">developmental plasticity<\/a><\/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 11.6: 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image10-1.jpg\" alt=\"Front view of black and white skull, missing lower jawbone.\" width=\"228\" height=\"228\" \/><figcaption class=\"wp-caption-text\">Figure 11.7: 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 11.8). 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 11.8: 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 11.5), 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\">It is theorized that 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. As discussed in Chapter 10, 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_1683_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 11.9). 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 11.9: 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 behaviour 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<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><\/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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image6-2.png\" alt=\"Lateral view of skull with large brow ridges.\" width=\"289\" height=\"289\" \/><figcaption class=\"wp-caption-text\">Figure 11.10: 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 11.10). 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_1683_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_1683_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_1683_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.<\/span><\/p>\n<div class=\"__UNKNOWN__\">\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 colour gene. Since chimpanzees have light skin under their hair, it is probable that early hominins also had light skin colour. 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<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 11.1). 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 11.11 and 11.12.<\/span><\/p>\n<figure style=\"width: 594px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 11.11:\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 11.12: 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 centre 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>\n<h2 class=\"__UNKNOWN__\"><span style=\"color: #000000\"><em>Homo erectus <\/em>Lifeways<\/span><\/h2>\n<div class=\"__UNKNOWN__\">\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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 11.13: 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_1683_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 11.13. 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<h3 class=\"import-Normal\"><span style=\"color: #000000\"><strong>Subsistence and Diet<\/strong><\/span><\/h3>\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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5-2.png\" alt=\"Five men excavating and note-taking at an archaeological site.\" width=\"253\" height=\"380\" \/><figcaption class=\"wp-caption-text\">Figure 11.14: 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 11.14), 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. 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, researchers propose that hominins began to cut up, tenderize, and process meat and plants, such that they did not have to use their teeth so vigorously.<\/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>It is proposed that the larger brain and body size of <em>Homo erectus<\/em> were fuelled 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. It is suggested that 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\u2014 might have given them greater flexibility in adapting to different environments. 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_1683_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_1683_1559\">human behavioural ecology<\/a><\/strong>, explored in more detail in Appendix C, examines the roots of human behaviour 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=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/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 11.15: 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 behavioural ecologists who study modern hunter-gatherer societies have observed that foraging is no easy business (Figure 11.15). 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_1683_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><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1560\">mosaic evolution<\/a><\/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 12), 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 behaviour 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 behavioural advancements, it now looks like many behaviours 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 behaviours; 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 behavioural 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 behaviours 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 behavioural ecology<\/strong>: The study of human behaviour 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<\/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>. Produced 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>. 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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. 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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\">Acknowledgements<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>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1683_1695\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1695\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<div class=\"textbox\">\n<p>Learn more about <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/about-the-authors\/\">authors and editors<\/a>,<a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/student-contributors\/\"> Hess' student contributions<\/a>, and <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/versioning-history\/\">versioning history<\/a>.<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\">Learning Objectives<\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Distinguish between adaptations and adjustments as ways of coping with environmental stressors.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Provide examples of adjustments humans use to cope with thermal stressors.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Describe how specific patterns of human adaptations and adjustments are correlated to natural selection processes.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Summarize the role of solar radiation in variations of human skin tone, and explain why reduced pigmentation is advantageous in northern latitudes.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Compare and contrast the various genetic mutations present in Tibetan and Ethiopian populations that allow them to survive at high altitudes.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Define the relationship between specific genetic mutations in some human populations and certain infectious diseases, such as the sickle-cell trait mutation and malarial infection.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p>As early humans left Africa and spread across the globe, they faced numerous challenges related to their new environments. Beyond genetically influenced changes in physiology as a result of evolution, humans have developed lifestyle strategies to cope with and even thrive in a wide range of habitats. The ways populations of humans met such challenges, coupled with their geographic separation throughout the majority of the last two hundred thousand years, have led to the many forms of adaptation in our species. This chapter focuses on the complexities of modern human variation through the lens of human evolutionary history.<\/p>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Stress and Homeostasis<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">All organisms, including humans, must maintain a baseline of normal functions within their cells, tissues, and organs to survive. This constancy of internal functions is referred to as <strong>homeostasis.<\/strong> Homeostatic regulation, however, may be disrupted by a variety of both external and internal stimuli known as <strong>stressors<\/strong>. Within limits, all organisms have evolved certain physiological mechanisms to respond to stressors in an effort to maintain homeostasis. The range of changes in the physiology (function), morphology (form), and\/or behaviour of organisms in response to their environments and potential stressors is regulated by its <strong>phenotypic<\/strong><strong> plasticity<\/strong>. Coping with these stressors led to the development of both <strong>adjustments<\/strong> (behavioural, acclimatory, and developmental) and <strong>adaptations<\/strong>, which are explained in detail in the following sections.<\/p>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Adjustments and Adaptations<\/h2>\n<h3 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Adjustments<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The term <em>adjustment<\/em> refers to an organism\u2019s nongenetic way of coping with the stressors of its environment. Although adjustments themselves are nongenetic in nature, the ability of an organism to experience or develop an adjustment is based on its phenotypic plasticity, which is linked to its evolutionarily guided genetic potential. Adjustments occur exclusively on the individual level. As such, different individuals within a population may experience a wide range of possible adjustments in response to a similar stressor. In general, the three main forms of adjustment are: behavioural, acclimatory, and developmental.<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><em>Behavioural Adjustments<\/em><\/h4>\n<figure style=\"width: 390px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2023\/06\/image3-3.jpg\" alt=\"Underwater photograph of scuba diver exploring the ocean floor.\" width=\"390\" height=\"293\" \/><figcaption class=\"wp-caption-text\">Figure 15.1: Notice the lack of full-spectrum colour in this photo of a deep-water diver as well as the diver\u2019s use of specialized equipment, such as a breathing apparatus to deliver gasses for respiration, a bodysuit to ensure thermal regulation, and a flashlight to increase visibility in the low-light setting. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-13\/\">Deep water diver<\/a> by Leslie E. Fitzpatrick 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\">When you are cold, do you reach for a blanket? When you are warm, do you seek out shelter cooled by an air-conditioning system? If so, you have likely been influenced to do so by the culture in which you were raised. As noted earlier in this textbook, the term <em>culture<\/em> refers to a collection of shared, learned beliefs and behaviours among individuals within a discrete population. <strong>Behavioural adjustments<\/strong> are regarded as cultural responses to environmental stressors. These adjustments are temporary in nature and, since they are nongenetic, must be constantly altered to meet novel situations posed by the environment. For example, divers are able to reach extraordinary depths (in excess of 300 metres below the surface) within the water through the use of a specialized mixture of gasses for breathing, an apparatus for the delivery of the gasses, protective clothing, and gear to increase visibility. The deeper a diver descends, the more atmospheric pressure the diver experiences, resulting in increased levels of potentially toxic byproducts of respiration within the body. In addition, with increased depth there is a decrease in the ambient temperature of the water and a decrease in the availability of light within the visible spectrum. Deep-water divers are well-versed in the environmental stressors of open waters and employ a variety of strategies based on behavioural adjustments to meet such demands. From wearing protective clothing to help maintain the body\u2019s core temperature to waiting at a specific depth for a prescribed period of time to facilitate the expulsion of nitrogen gas that may have accumulated within the bloodstream, divers employ numerous behavioural adjustments to ensure their safety (Figure 15.1). Without these culturally mediated behavioural adjustments, a deep-water diver\u2019s first dive would also be their last.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In many developing countries, the use of refrigeration for the storage of perishable food products is uncommon; therefore, individuals within these cultures have developed a variety of behavioural adjustment strategies related to food preparation to address possible food spoilage. Through a cross-cultural analysis of spice use in recipes, Paul Sherman and Jennifer Billing (1999) determined that cultures closest to the equator, where temperatures are hotter, tend to use both a greater number and a wider variety of plant-based spices with bacteria-inhibiting phytochemical properties (e.g., garlic and onion). Antimicrobial properties of spices permits the consumption of foods, particularly animal-based protein sources, for a period of time beyond that which would be considered safe. There are some acclimatory adjustment benefits to the use of some pungent spices as well, which are explored in the following section.<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><em>Acclimatory Adjustments: Thermal Stressors<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Acclimatory adjustments<\/strong> are temporary, reversible changes in an organism\u2019s physiology in response to environmental stressors. Although they are not genetically determined, the range of acclimatory adjustments that an organism is capable of producing is linked to its underlying phenotypic plasticity and the duration and severity of the stressor. A good example of this is the human response to varying ambient temperatures.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">To understand human adjustments, we must first understand the thermodynamic mechanisms through which heat may be gained or lost. The four pathways for this are conduction, convection, evaporation, and radiation (Figure 15.2).<\/p>\n<figure style=\"width: 376px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image11-5.png\" alt=\"Person facing wall with arrows illustrating evaporation, conduction, radiation, and convection.\" width=\"376\" height=\"426\" \/><figcaption class=\"wp-caption-text\">Figure 15.2: Various thermodynamic mechanisms related to heat gain and loss in the human body. This process is decribed in the text below. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-13\/\">Mechanisms of heat transfer (Figure 14.2)<\/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 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Through <strong>conduction<\/strong> processes, heat will move from a warmer body to a cooler one through direct contact. An example of this is when you accidentally touch a hot cooktop with your hand and the heat is transferred from the cooktop to your skin.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">With <strong>convection<\/strong>, when a warm body is surrounded by a cooler fluid (e.g., air or water), heat will be transferred from the warmer body to the cooler fluid. This is why we will often employ the behavioural adjustment of wearing multiple layers of clothing during the winter in an effort to prevent heat loss to the cooler atmosphere. Conversely, if your body temperature is cooler than that of the air surrounding you, your body will absorb heat.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Depending on your physical condition, most people will begin to sweat around 37.2\u2103 to 37.7\u2103 (98.9\u2109\u201399.9\u2109). Sweating is an example of <strong>evaporation<\/strong>, which occurs when a liquid, such as the water within our bodies, is converted to a gas. Phase conversions, such as those underlying the evaporative processes of transforming liquids to gasses, require energy. In evaporation, this energy is in the form of heat, and the effect is to cool the body.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The final mechanism for heat loss within the human body is <strong>radiation<\/strong>, through which energy in the form of electromagnetic waves is produced at a wavelength that typically lies below that which is visible to the human eye. Although humans gain and lose heat from their bodies through radiation, this form of heat transfer is not visible. Humans are capable of losing and gaining heat through conduction, convection, and radiation; however, heat may not be gained through evaporation.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">As the ambient temperature decreases, it becomes increasingly difficult for the human body to regulate its core temperature, which is central to the maintenance of homeostasis. When an individual\u2019s body temperature falls below 34.4\u2103 (93.9\u00b0F), the brain\u2019s <strong>hypothalamus<\/strong> becomes impaired, leading to issues with body temperature control. A total loss of the ability to regulate body temperature occurs around 29.4\u2103 (84.9\u00b0F), which may result in death. When the ambient temperature falls below the critical temperature of 31\u2103 (87.8\u00b0F), a nude human body that is at rest will respond with a series of physiological changes to preserve homeostasis (Figure 15.3).<\/p>\n<figure style=\"width: 585px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image26-2.jpg\" alt=\"Body outlines illustrate differences in heat maintenance for different ambient environments.\" width=\"585\" height=\"347\" \/><figcaption class=\"wp-caption-text\">Figure 15.3: Example of overall body heat maintenance in cold and warm ambient environments. <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__-13\/\">Body heat maintenance in cold and warm (Figure 14.3)<\/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=\"margin-left: 0pt;text-indent: 0pt\">The human body experiences two main types of physiological responses to colder temperatures: those that increase the production of heat and those that seek to retain heat. The production of heat within the body is accomplished through short-term increases in the body\u2019s basal metabolic rate, such as shivering to increase muscular metabolism. An organism\u2019s basic metabolic rate is a measure of the energy required to maintain necessary body processes when the organism is at rest. Increases in basal metabolic rates, such as when we shiver from the cold, require increased consumption of energy-providing nutrients. Of course, such increases in metabolic rates are not infinite, as we may only consume a finite amount of nutrients. As with all <strong>acclimatory adjustments<\/strong>, an increase in the basal metabolic rate is merely temporary.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Of the physiological mechanisms to preserve heat already in the body, the most notable is <strong>vasoconstriction<\/strong>, or the constriction of peripheral capillaries in the skin. The decreased surface area of the capillaries through vasoconstriction results in less heat reaching the surface of the skin where it would be dissipated into the atmosphere. Vasoconstriction also leads to the maintenance of heat near the core of the body where the vital organs are located. As a trade-off, though, individuals are more at risk of cold-related injuries, such as frost-bite, which can lead to tissue necrosis (tissue death) in regions of the body that are most distant from the core (e.g., fingers, toes, nose, ears, cheeks, chin, etc.).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Just as cold stress presents challenges to maintaining homeostasis, heat does as well. In hot climates, the body will absorb heat from its surroundings (through conduction, convection, and radiation), resulting in potential heat-related disorders, such as heat exhaustion. When the human body is exposed to ambient temperatures above 35\u2103 (95\u00b0F), excess body heat will be lost primarily through evaporative processes, specifically through sweating. All humans, regardless of their environment, have approximately the same number of sweat glands within their bodies. Over time, individuals living in hot, arid environments will develop more sensitive forms of sweat glands resulting in the production of greater quantities of sweat (Best, Lieberman, and Kamilar 2019; Pontzer Et al. 2021). In an effort to prevent dehydration due to this form of acclimatory adjustment, there will be an additional reduction in the volume of urine produced by the individual (Pontzer Et al. 2021).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">As noted in the previous section, some cultural groups, particularly those in equatorial regions, add pungent spices to their foods to inhibit the colonization of bacteria (Sherman and Billing 1999). Although adding spices to decrease spoilage rates is a behavioural adjustment, the application of some forms of peppers triggers an acclimatory adjustment as well. Compounds referred to as capsaicinoids are the secondary byproducts of chili pepper plants\u2019 metabolism and are produced to deter their consumption by some forms of fungi and mammals. When mammals, such as humans, consume the capsaicinoids from chili peppers, a burning sensation may occur within their mouths and along their digestive tracts. This burning sensation is the result of the activation of capsaicin receptors along the body\u2019s nerve pathways. Although the peppers themselves may be at ambient temperature so their consumption is not causing any form of body temperature increase, the human body perceives the pepper as elevating its core temperature due to the activation of the capsaicin receptors. This causes the hypothalamus to react, initiating sweating in an attempt to lower body temperature and maintain homeostasis. The increased piquancy (application of pungent spices to food) as a means of inhibiting food-borne bacterial colonization in warm climates, as well as spices\u2019 ability to trigger sweating processes as a method for cooling the body, is an example of the intersection between behavioural and acclimatory adjustments that utilized within certain populations.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In addition to increased sweat production to maintain homeostasis in excess heat, <strong>vasodilation<\/strong> may occur (Figure 15.4). Vasodilation is an expansion of the capillaries within the skin leading to a more effective transfer of heat from within the body to the exterior to allow conductive, convective, radiative, and evaporative (sweating) processes to occur.<\/p>\n<figure style=\"width: 499px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image27-4.png\" alt=\"Side-by-side sketches of capillaries under the skin for heat and cold.\" width=\"499\" height=\"445\" \/><figcaption class=\"wp-caption-text\">Figure 15.4: The vasoconstriction processes occur within the peripheral vascular system when an individual is exposed to cold ambient temperatures and the vasodilation that occurs in warmer environments. <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__-13\/\">Vasoconstriction and vasodilation (Figure 14.4)<\/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\">Physiologically based acclimatory adjustments to hot, dry climates may be complemented by behavioural adjustments as well. For example, individuals in such climates may limit their physical activity during the times of day when the temperature is typically the hottest. Additionally, these individuals may wear loose-fitting clothing that covers much of their skin. The looseness of the clothing allows for air to flow between the clothing and the skin to permit the effective evaporation of sweat. Although it may seem counterintuitive to cover one\u2019s body completely in a hot climate, the covering of the skin keeps the sun\u2019s rays from directly penetrating the skin and elevating the body\u2019s core temperature.<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><em>Acclimatory Adjustments: Altitudinal Stressors<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The challenges posed by thermal conditions are but one form of environmental stressor humans must face. High-altitude environments, which are defined as altitudes in excess of 2,400 metres above sea level (masl) or 7,874 feet above sea level (fasl), pose additional challenges to the maintenance of homeostasis in humans. Some of the main stressors encountered by those living within high-altitude environments include decreased oxygen availability, cold temperatures, low humidity, high wind speed, a reduced nutritional base, and increased solar radiation levels. Of these challenges, the most significant is the decreased availability of oxygen.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">To visualize how altitude affects the availability of oxygen, imagine two balloons that are each filled with the same quantity of oxygen molecules. One of these balloons is positioned at sea-level and the other is placed high upon a mountain peak. For the balloon at sea level, there is more atmospheric pressure pressing down on the molecules within this balloon. This leads to the oxygen molecules within the sea level balloon being forced into a more compact organization. In contrast, the mountain peak balloon has less atmospheric pressure pressing down on it. This leads to the oxygen molecules within that balloon spreading out from each other since they are not being forced together quite as strongly. This example highlights the availability of oxygen molecules in each breath than we take in low- versus high-altitude environments. At 5,500 masl (approximately 18,000 fasl), the atmospheric pressure is approximately 50% of its value at sea level (Peacock 1998). At the peak of Mount Everest (8,900 masl or approximately 29,200 fasl), the atmospheric pressure is equivalent to only about 30% of their sea level amounts (Peacock 1998; Figure 15.5).<\/p>\n<figure style=\"width: 482px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image21-5.png\" alt=\"Atmospheric pressure decreases in density as a person climbs a mountain and increases in altitude. \" width=\"482\" height=\"418\" \/><figcaption class=\"wp-caption-text\">Figure 15.5: As altitude increases, atmospheric pressure decreases, which allows for more space between air molecules. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-13\/\">Atmospheric pressure (Figure 14.5)<\/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\">Due to decreased availability of oxygen at higher altitudes, certain acclimatory adjustments are required to ensure the maintenance of homeostasis for individuals other than those who were gestated, born, and raised at high altitude. For these people, their rate of breathing will increase to permit greater quantities of air containing oxygen into the lungs when they ascend into higher altitude environments. An increased speed and depth of breathing, which is referred to as <strong>hyperpnea<\/strong>, is not sustainable indefinitely; thus, the rate of breathing begins to decrease as the person becomes acclimatized to the altitude. During the initial phases of high-altitude-related hyperpnea, the heart begins to beat faster but the amount of blood pushed through during each beat decreases slightly. In addition, the body will divert energy from noncritical bodily functions, such as digestive processes.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Once the atmospheric oxygen reaches the alveoli (small air sacs) in the lungs, it spreads across the alveolar membrane and enters <strong>erythrocytes<\/strong> (red blood cells). As oxygen reaches the alevoli\u2019s erythrocytes, it loosely binds with hemoglobin (an iron-rich protein) contained in the erythrocytes. When the erythrocytes carrying the hemoglobin-bound oxygen molecules reach capillaries where the partial pressure of oxygen is relatively low, oxygen will be released by the hemoglobin so that it is free for diffusion into body cells. Similar to acclimatory adjustments related to thermal conditions (e.g., shivering or sweating), those related to high altitude may not be infinitely sustained due to their energetically expensive nature.<\/p>\n<figure style=\"width: 430px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image24-3.jpg\" alt=\"Newborn lays face down on hospital bedding. Photograph is blue indicating UV light.\" width=\"430\" height=\"323\" \/><figcaption class=\"wp-caption-text\">Figure 15.6: Premature infant born at 30 weeks, 4 days gestation to a mother with altitudinal-induced preeclampsia. Blue light assists the infant\u2019s liver with processing high levels of bilirubin. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-13\/\">Premature infant<\/a> by Leslie E. Fitzpatrick 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\">Although the long-term acclimatory adjustments that an individual from low altitude experiences in a high-altitude environment may permit them to reside there successfully, reproduction within such settings is frequently complicated. With increased altitude comes an increased risk of miscarriage, lower birth weights, and higher infant mortality rates. As the pregnant person\u2019s body seeks to preserve its own homeostasis, there is often a decreased rate and volume of blood flow to the uterus as compared to a pregnant person of similar physiological condition at a lower altitude (Moore, Niermeyer, and Zamudio 1998). This results in a decrease in the amount of oxygen that will be passed through the uterus and placenta to the developing fetus. In addition, pregnant people who experience pregnancy at higher altitudes are more prone to developing preeclampsia (severe elevation of blood pressure), which is linked to increased rates of both fetal and maternal death (Moore, Niermeyer, and Zamudio 1998; Figure 15.6).<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><em>Developmental Adjustments <\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Developmental adjustments occur only in individuals who spent their developmental period (i.e., childhood and adolescence) within a high-altitude environment; they do not apply to those who moved into these environments in the post developmental (i.e., adult) phase. Furthermore, the degree of developmental adjustment within an individual is directly related to their underlying phenotypic plasticity as well as the amount of time during the crucial growth and development period that the individual resides within the challenging environment. Although humans have the remarkable capacity to develop and survive within environments that are not overly conducive to the successful maintenance of homeostasis, there are definitely physiological costs associated with this ability.<\/p>\n<figure style=\"width: 369px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image18-2.jpg\" alt=\"Two people walking down a street wearing brightly colored woven traditional clothing.\" width=\"369\" height=\"277\" \/><figcaption class=\"wp-caption-text\">Figure 15.7: Two individuals from a high-altitude region of the Peruvian Andes. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Andahuaylas_Peru-_two_women_walking_down_street.jpg\">Andahuaylas Peru-two women walking down street<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:ThayneT\">Thayne Tuason<\/a> has been modified (cropped) and is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/deed.en\">CC BY 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In general, high-altitude natives tend to grow more slowly and physically mature later than their low-altitude counterparts (Figure 15.7). Lowered growth and maturity rates are linked not only to the increased physiological demands placed on the body due to the decreased partial pressure of oxygen but also to reductions in the quality of the nutritional base at higher altitudes. Increased terrain complexity, elevated solar radiation levels, and higher wind speeds coupled with the lower temperatures and humidity levels found at high altitudes leads to difficulties with growing and maintaining crops and raising livestock. Overall, as altitude rises, the quality of the available nutritional base goes down, which is correlated to a lack of the nutrients necessary to ensure proper physiological growth and development in humans. Thus, even though individuals may be able to develop and grow within high-altitude environments, they may not reach their full genetically mediated growth potential as they would in a lower-altitude environment.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Not all developmental adjustments are linked to environmental pressures such as climate or altitude; rather, some of these adjustments are correlated to sociocultural or behavioural practices. Some of these adjustments may affect the physiological appearance of an individual when they are practiced consistently during the development and growth phases.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Sudden infant death syndrome (SIDS) has no definitive cause; however, the American Academy of Pediatrics published a report in 1992 linking SIDS to infants (under the age of one) sleeping on their stomachs. The \u201cBack to Sleep\u201d campaign championed by the American Academy of Pediatrics helped educate members of the medical community as well as the public that the best sleep position for infants is on their backs (American Academy of Pediatrics 2000).<\/p>\n<\/div>\n<figure id=\"attachment_420-3\" aria-describedby=\"caption-attachment-420-3\" style=\"width: 562px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-419\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/14.8.jpg\" alt=\"Brachycephaly drawings show a flattened, symmetrical head. Plagiocephaly shows asymmetrical flattening of the skull.\" width=\"562\" height=\"219\" \/><figcaption id=\"caption-attachment-420-3\" class=\"wp-caption-text\">Figure 15.8: These sketches illustrate a top and side view of brachycephaly (left and middle images, respectively) and plagiocephaly (right image). Credit: Brachycephaly and plagiocephaly original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) 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 class=\"__UNKNOWN__\">\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Placing infants on their backs to sleep has led to decreased infant mortality (death) rates due to SIDS; however, it has led to an unintended consequence: infant cranial deformation. The cranial deformations experienced by infants who sleep solely on their back tend to manifest in one of two forms: brachycephaly and plagiocephaly (Roby Et al. 2012; Figure 15.8). With positional brachycephaly, the back of the infant\u2019s head appears rather uniformly flattened due to repetitive contact with a flat surface, such as a crib mattress or car seat back. In cases of positional plagiocephaly, the back of the infant\u2019s head appears asymmetrically flattened. This asymmetry is typically due to an uneven distribution of mechanical forces resulting from the manner in which the infant\u2019s head is in contact with a flat surface. The forms of cranial deformation resulting from sleep positioning do not affect the infant\u2019s brain development. For many individuals, the appearance of the deformation is minimized during later development. Still, some individuals will maintain the pattern of cranial deformation acquired during their infancy throughout their lives. The unintentional cranial deformation resulting from placing infants on their backs to sleep as a means of preventing SIDS-related deaths is a physiological indicator of a behavioural adjustment.<\/p>\n<\/div>\n<h3 class=\"__UNKNOWN__\"><strong>Adaptations<\/strong><\/h3>\n<div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">As we have just explored, survival and reproduction at high altitudes present numerous physiological challenges for most humans. The behavioural, acclimatory, and developmental adjustments discussed above are all related to the phenotypic plasticity of the individual; however, most adjustments are temporary in nature and they affect a single individual rather than all individuals within a population. But what if the physiological changes were permanent? What if they affected all members of a population rather than just a single individual? The long-term, microevolutionary (i.e., genetic) changes that occur within a population in response to an environmental stressor are referred to as an adaptation. From an evolutionary standpoint, the term <em>adaptation<\/em> refers to a phenotypic trait (i.e., physiological\/morphological feature or behaviour) that has been acted upon by natural selection processes to increase a species\u2019 ability to survive and reproduce within a specific environment. Within the field of physiology, the term <em>adaptation<\/em> refers to traits that serve to restore homeostasis. The physiology-based interpretation of adaptations presumes that all traits serve a purpose and that all adaptations are beneficial in nature; however, this may be a fallacy, since some traits may be present without clear evidence as to their purpose. As such, during the following discussion of various forms of adaptations in human populations, we will focus our attention on phenotypic traits with an evidence-based purpose.<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><em>Adaptation: <\/em><em>Altitudinal Adaptation<\/em><\/h4>\n<\/div>\n<figure id=\"attachment_420-4\" aria-describedby=\"caption-attachment-420-4\" style=\"width: 619px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-420\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/14.9.jpg\" alt=\"The Simian Plateau in northeast Africa and Tibetan Plateau in southern Asia.\" width=\"619\" height=\"339\" \/><figcaption id=\"caption-attachment-420-4\" class=\"wp-caption-text\">Figure 15.9: Highlighted regions feature (from left to right) the Simian (Ethiopian) and Tibetan Plateau high-altitude regions. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:World_Map_of_HVR_adaptation_in_high_altitude_populations.jpg\">World Map of HVR adaptation in high altitude populations<\/a> by Chkuu has been modified (cropped, Andean region colour change) and is under <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<div class=\"__UNKNOWN__\">\n<p>As mentioned in the previous section, there is genomic research supporting the evolutionary selection of certain phenotypes and their corresponding genotypes within indigenous high-altitude populations across the globe. The following discussion focuses on two high-altitude indigenous populations from Tibet and Ethiopia (Figure 15.9). Although these populations share many common genetic traits based on relatively similar evolutionary histories influenced by similar environmental stressors, there is support for local genetically based adaptation as well, based on different genes being acted upon by environmental stressors that may be unique to Tibet and Ethiopia (Bigham 2016).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Tibetan populations have resided in the Tibetan Plateau and Himalayan Mountain regions at elevations exceeding 4,000 masl (13,100 fasl) for at least the past 7,400 years (Meyer Et al. 2017). There is evidence of a genetic exchange event involving Tibetan populations and Denisovans around 48,700 years ago, which introduced a haplogroup involving mutations of the <em>EPAS1<\/em> gene (Zhang Et al. 2021). The <em>EPAS1<\/em> is involved in the regulation of erythrocytes and hemoglobin. For individuals originating in lower-altitude environments, <em>EPAS1 <\/em>stimulates increased erythrocyte production in high-altitude environments as a temporary acclimatory adjustment. For indigenous high-altitude populations of Tibet, the <em>EPAS1<\/em> gene mutation introduced by Denisovan introgression inhibits increased erythrocyte production, which reduces potential negative effects (e.g., stroke or heart attack) associated with long-term high levels of erythrocyte production (Gray Et al. 2022; Zhang Et al. 2021). The erythrocyte count of high-altitude Tibetans with the <em>EPAS1<\/em> point mutation is about the same as for individuals residing at sea level.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Populations indigenous to the Semien Plateau of Ethiopia, such as the Oromo and Amhara, share a similar but not identical <em>EPAS1<\/em> point mutation with the Tibetan population (Bigham 2016); however, there is no indication that this mutation was derived from Denisovan introgression. The <em>EPAS1<\/em> mutations occurred independently from each other; however, their effects are still similar in that they permit the Tibetan and Ethiopian populations to survive at high altitudes. Not all adaptations are related to life in high-altitude environments, however. In the following sections, we will address two more general examples of adaptation in human populations: variations in skin colour and differences in body build.<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><em>Adaptation: Skin Tone<\/em><em> Basics<\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">When you think about your own skin tone and compare it to members of your family, do you all possess exactly the same shade? Are some members of your family darker than others? What about your friends? Your classmates? Skin tone occurs along a continuum, which is a reflection of the complex evolutionary history of our species. The expression of skin tone is regulated primarily by melanin and hemoglobin. <strong>Melanin<\/strong> is a dark brown-black pigment that is produced by the oxidation of certain amino acids (e.g., tyrosine, cysteine, phenylalanine) in melanocytes. <strong>Melanocytes<\/strong> are specialized cells located in the base layer (stratum basale) of the skin\u2019s epidermis as well as several other areas within the body (Figure 15.10). Within the melanocytes, melanin is produced in the special organelle called a melanosome. Melanosomes serve as sites for the synthesis, storage, and transportation of melanin. Melanosomes transport the melanin particles through cellular projections to epidermal skin cells (keratinocytes) as well as to the base of the growing hair root. In the eye, however, melanin particles produced by the melanosomes remain present within the iris and are not transported beyond their origin location. The two main forms of melanin related to skin, hair, and eye colour are eumelanin and pheomelanin. All humans contain both eumelanin and pheomelanin within their bodies; however, the relative expression of these two forms of melanin determines an individual\u2019s overall colouring. Eumelanin is a brown-to-black coloured melanin particle while pheomelanin is more pink-to-red coloured. Individuals with darker skin or hair colour have a greater expression of eumelanin than those with lighter-coloured skin and blonde or red hair.<\/p>\n<figure style=\"width: 553px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image12-2.jpg\" alt=\"Melanocytes and melanosomes are compared from light and dark skin tones.\" width=\"553\" height=\"436\" \/><figcaption class=\"wp-caption-text\">Figure 15.10: Diagram featuring the relative numbers of melanocytes and melanosomes in light and dark shades of skin tone. <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:\/\/openstax.org\/books\/anatomy-and-physiology-2e\/pages\/5-1-layers-of-the-skin#fig-ch05_01_07\">Skin Pigmentation (Anatomy and Physiology, Figure 5.8)<\/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<h4 class=\"import-Normal\"><em>Adaptation<\/em><em>: <\/em><em>Melanogenesis <\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Although all humans have approximately the same number of melanocytes within their epidermis, the production of melanin by these melanocytes varies. There are two forms of melanogenesis (the process through which melanocytes generate melanin): basal and activated. As discussed previously, the expression of eumelanin and pheomelanin by the melanocytes is genetically regulated through the expression of specific receptors (e.g., <em>MC1R<\/em>) or other melanocyte components (e.g., <em>MFSD12<\/em>)<em>.<\/em> <strong>Basal melanogenesis <\/strong>is dependent upon an individual\u2019s inherent genetic composition and is not influenced by external factors. <strong>Activated melanogenesis<\/strong> occurs in response to ultraviolet radiation (UV) exposure, specifically UV-B (short UV wave) exposure. Increased melanogenesis in response to UV-B exposure serves to provide protection to the skin\u2019s innermost layer called the hypodermis, which lies below the epidermis and dermis (Figure 15.11). Melanin in the skin, specifically eumelanin, effectively absorbs UV-B radiation from light\u2014meaning that it will not reach the hypodermal layer. This effect is often more apparent during periods of the year when people tend to be outside more and the weather is warmer, which leads to most donning fewer protective garments. The exposure of skin to sunlight is, of course, culturally mediated with some cultures encouraging the covering of skin at all times.<\/p>\n<figure style=\"width: 395px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image10-6.png\" alt=\"Cross-section of skin illustrating UVA rays penetrate deeper than UVB. \" width=\"395\" height=\"407\" \/><figcaption class=\"wp-caption-text\">Figure 15.11: Penetration of skin layers by UVA and UVB rays. UVB rays penetrate only through the epidermis. UVA rays penetrate much deeper, through the dermis. Credit: <a href=\"https:\/\/cnx.org\/contents\/FPtK1zmh@16.7:RxywCGkA@10\/5-1-Layers-of-the-Skin\">Skin Pigmentation (Anatomy and Physiology, Figure 5.8)<\/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\">As previously noted in this chapter, hemoglobin is an iron-rich protein that binds with oxygen in the bloodstream. For individuals with lighter-coloured skin, blood vessels near the surface of the skin and the hemoglobin contained within those vessels is more apparent than in individuals with darker skin. The visible presence of hemoglobin coupled with the pink-to-red tone of the pheomelanin leads to lighter-skinned individuals having a pale pink skin tone. Individuals with lighter skin more readily absorb UV radiation as their basal melanin expression is directed more toward the production of pheomelanin than eumelanin. But why are there so many variations in skin tone in humans? To answer this question, we now turn toward an exploration of an evolutionary-based adaptation of skin tone as a function of the environment.<\/p>\n<h4 class=\"import-Normal\"><em>Adaptation: <\/em><em>Evolutionary Basis for Skin Tone Variation<\/em><\/h4>\n<figure style=\"width: 337px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image22-4.png\" alt=\"Table discussing hair and skin, folate and UV rays, leaving Africa, vitamin D, and selective pressures.\" width=\"337\" height=\"843\" \/><figcaption class=\"wp-caption-text\">Figure 15.12: Evolutionary basis for human skin colour variation. <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__-13\/\">Evolutionary basis for human skin color variation (Figure 14.12)<\/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>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Skin cancer is a significant concern for many individuals with light skin tone as the cumulative exposure of the epidermis and underlying skin tissues to UV radiation may result in the development of abnormal cells within those tissues, leading to malignancies. Although darker-skinned individuals are at risk for skin cancer as well, they are less likely to develop it due to increased levels of melanin, specifically eumelanin, in their skin. Even though skin cancer is a serious health concern for some individuals, most skin cancers occur in the postreproductive years; therefore, it is improbable that evolutionary forces favouring varying melanin expression levels are related to a selective pressure to avoid such cancers. Furthermore, if avoiding skin cancer were the primary factor driving the evolution of various skin tones, then it reasons that everyone would have the most significant expression of eumelanin possible. So, why do we have different skin tones (Figure 15.12)?<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The term <strong>cline<\/strong> (introduced in Chapter 14) refers to the continuum or spectrum of gradations (i.e., levels or degrees) from one extreme to another. With respect to skin tone, the various tonal shades occur clinally with darker skin being more prevalent near the equator and gradually decreasing in tone (i.e., decreased melanin production) in more distant latitudes. For individuals who are indigenous to equatorial regions, the increased levels of melanin within their skin provides them with a measure of protection against both sunburn and sunstroke because the melanin is more reflective of UV radiation than hemoglobin. In cases of severe sunburn, eccrine glands are affected, resulting in an individual\u2019s ability to sweat being compromised. As sweat is the body\u2019s most effective means of reducing its core temperature to maintain homeostasis, damage to the eccrine glands may lead to numerous physiological issues related to heat that may ultimately result in death.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Even though avoiding severe sunburn and sunstroke is of great importance to individuals within equatorial regions, this is likely not the primary factor driving the evolutionary selection of darker skin within these regions. It has been proposed that UV radiation\u2019s destruction of <strong>folic acid<\/strong>, which is a form of B-complex vitamin, may have led to the selection of darker skin in equatorial regions. For pregnant people, low levels of folic acid within the body during gestation may lead to defects in the formation of the brain and spinal cord of the fetus. This condition, which is referred to as spina bifida (Figure 15.13), often significantly reduces an infant\u2019s chances of survival without medical intervention. In people producing sperm, low levels of folic acid within the body reduce sperm quantity and quality. Thus, in geographic regions with high UV radiation levels (i.e., equatorial regions), there appears to be an evolutionarily driven correlation between darker skin and fertility.<\/p>\n<figure style=\"width: 364px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image19-2-1.jpg\" alt=\"Infant with a bulging spinal cord above the lower spine.\" width=\"364\" height=\"294\" \/><figcaption class=\"wp-caption-text\">Figure 15.13: Infant with open neural tube defect in lower (lumbar) region of the spine (right). A close-up of the open neural tube defect within the spinal column (left) shows the dura matter, which is ordinarily protected within the spine, is exposed on the surface of the skin. The spinal cord sits near the surface, when it should be protected within the spinal column. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Spina-bifida.jpg\">Spina-bifida<\/a> by the <a href=\"https:\/\/www.cdc.gov\/\">Centers for Disease Control and Prevention<\/a> is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p>If darker skin tone is potentially correlated to more successful reproduction, then why do lighter shades of skin exist? One hypothesis is that there is a relationship between lighter skin tone and vitamin D synthesis within the body. When skin is exposed to the UV-B radiation waves in sunlight, a series of chemical reactions occur within the epidermis leading to the production of vitamin D3, which is a fat-soluble vitamin that assists the body with absorbing calcium and phosphorus in the small intestine. These nutrients are among those that are critical for the proper growth and maintenance of bone tissue within the body. In the absence of adequate minerals, particularly calcium, bone structure and strength will be compromised, leading to the development of rickets during the growth phase. Rickets is a disease affecting children during their growth phase. It is characterized by inadequately calcified bones that are softer and more flexible than normal. Individuals with rickets will develop a true bowing of their legs, which may affect their mobility (Figure 15.14). In addition, deformation of pelvic bones in people who may become pregnant may occur as a result of rickets, leading to complications with reproduction. In adults, a deficiency in vitamin D3 will often result in osteomalacia, which is a general softening of the bones due to inadequate mineralization. As noted, a variety of maladies may occur due to the inadequate production or absorption of vitamin D3, as well as the destruction of folate within the human body. Therefore, from an evolutionary perspective, natural selection should favour a skin tone that is best suited to a given environment.<\/p>\n<figure style=\"width: 263px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image9-2-1.jpg\" alt=\"Historic photo of three young children each with visible lower limb curvature.\" width=\"263\" height=\"315\" \/><figcaption class=\"wp-caption-text\">Figure 15.14: Children with rickets in various developmental stages. Credit: <a href=\"https:\/\/wellcomecollection.org\/works\/m3eu5snb\">Rachitis, stages of development for children<\/a> [slide numbers 7181 and 7182; photo number: M0003399] by <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>In general, the trend related to lighter skin pigmentation further from the equator follows a principle called <strong>Gloger\u2019s Rule<\/strong>. This rule states that within the same species of mammals the more heavily pigmented individuals tend to originate near the equator while lighter-pigmented members of the species will be found in regions further from the equator. Gloger\u2019s Rule applies latitudinally; however, it does not appear to hold for certain human populations near the poles. Specifically, it does not apply to the Inuit people (Figure 15.15), who are indigenous to regions near the North Pole and currently reside in portions of Canada, Greenland, Alaska, and Denmark. The Inuit have a darker skin tone that would not be anticipated under the provisions of Gloger\u2019s Rule. The high reflectivity of light off of snow and ice, which is common in polar regions, necessitates the darker skin tone of these individuals to prevent folic acid degradation just as it does for individuals within equatorial regions. The consumption of vitamin D\u2013rich foods, such as raw fish, permits the Inuit to reside at high latitudes with darker skin tone while preventing rickets.<\/p>\n<h4 class=\"import-Normal\"><em>Adaptation: <\/em><em>Shape and Size Variations<\/em><\/h4>\n<figure style=\"width: 353px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image6-3-1.jpg\" alt=\"Historic photo of a person carrying a very large fish laid on a pack.\" width=\"353\" height=\"296\" \/><figcaption class=\"wp-caption-text\">Figure 15.15: Copper Inuk (from Bernard Harbour, Nunavut) with lake trout on his back (1915). Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Copper_Inuk_with_lake_trout_on_his_back_near_Bernard_Harbour_(51007).jpg\">Copper Inuk with lake trout on his back near Bernard Harbour, Northwest Territories (Nunavut)<\/a> (1915) by George H. Wilkins is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0 License<\/a>. This image is part of the Photographic Archives of the Canadian Museum of History.<\/figcaption><\/figure>\n<p>In addition to natural selection playing a role in the determination of melanin expression, it plays a significant role in the determination of the shape and size of the human body. As previously discussed, the most significant thermodynamic mechanism of heat loss from the body is radiation. At temperatures below 20\u2103 (68\u2109), the human body loses around 65% of its heat to radiative processes; however, the efficiency of radiation is correlated to the overall body shape and size of the individual. There is a direct correlation between the ratio of an object\u2019s surface area to mass and the amount of heat that may be lost through radiation. For example, two objects of identical composition and mass are heated to the same temperature. One object is a cube and the other is a sphere. Which object will cool the fastest? Geometrically, a sphere has the smallest surface area per unit mass of any three-dimensional object, so the sphere will cool more slowly than the cube. In other words, the smaller the ratio of the surface area to mass an object has, the more it will retain heat. With respect to the cube in our example, mass increases by the cube, but surface area may increase only by the square, so size will affect the mass to surface area ratio. This, in general, holds true for humans, as well.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In regions where temperatures are consistently cold, the body shape and size of individuals indigenous to the area tend to be more compact. These individuals have a relatively higher body mass to surface area (i.e., skin) than their counterparts from equatorial regions where the average temperatures are considerably warmer. Individuals from hot climates, such as the Fulani (Figure 15.16) of West Africa, have limbs that are considerably longer than those of individuals from cold climates, such as the Inuit of Greenland (Figure 15.16). Evolutionarily, the longer limbs of individuals from equatorial regions (e.g., the Fulani) provide a greater surface area (i.e., lower body mass to surface area ratio) for the dissipation of heat through radiative processes. In contrast, the relatively short limbs of Arctic-dwelling people, such as the Inuit, allows for the retention of heat because there is a decreased surface area through which heat may radiate away from the body.<\/p>\n<figure style=\"width: 385px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image20-2.jpg\" alt=\"Person on left is stocky, person on right has a narrow body breadth.\" width=\"385\" height=\"570\" \/><figcaption class=\"wp-caption-text\">Figure 15.16: The individual on the left is typical of one adapted to a cold environment where the conservation of heat in the body\u2019s core is of critical importance.The individual on the right could be adapted to a tropical environment where the rapid dispersal of heat is necessary to maintain homeostasis. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Greenland_1999_(01).jpg\">Greenland 1999 (01)<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Vadeve\">Vadeve<\/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\">As described above, there are certain trends related to the general shape and size of human bodies in relation to the thermal conditions. To better describe these trends, we turn to a couple of general principles that are applicable to a variety of species beyond humans. <strong>Bergmann\u2019s Rule<\/strong> predicts that as average environmental temperature decreases, populations are expected to exhibit an increase in weight and a decrease in surface area (Figure 15.17a). Also, within the same species of homeothermic animals, the relative length of projecting body parts (e.g., nose, ears, and limbs) increases in relation to the average environmental temperature (Figure 15.17b). This principle, referred to as <strong>Allen\u2019s Rule<\/strong>, notes that longer, thinner limbs are advantageous for the radiation of excess heat in hot environments and shorter, stockier limbs assist with the preservation of body heat in cold climates. A measure of the crural index (crural index = tibia length <span style=\"font-size: NaNpt;color: #;text-decoration: none\">\u00f7 <\/span>femur length) of individuals from various human populations provides support for Allen\u2019s Rule since this value is lower in individuals from colder climates than it is for those from hot climates. The crural indices for human populations vary directly with temperature, so individuals with higher crural index values are generally from regions with a warmer average environmental temperature. Conversely, the crural indices are lower for individuals from regions where there are colder average temperatures.<\/p>\n<figure style=\"width: 533px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image16-6.png\" alt=\"Illustration of an elk (left) and gazelle (right).\" width=\"533\" height=\"319\" \/><figcaption class=\"wp-caption-text\">Figure 15.17a: These organisms are representative of Bergmann\u2019s rule. The animal on the left depicts an ungulate from a cooler environment with increased body weight and decreased surface area, compared to the slender ungulate on the right. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-13\/\">Bergmann\u2019s Rule (Figure 14.16a)<\/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<figure style=\"width: 521px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image14-4.png\" alt=\"Three types of rabbits with different ear and forelimb lengths.\" width=\"521\" height=\"264\" \/><figcaption class=\"wp-caption-text\">Figure 15.17b: These animals are representative of Allen\u2019s rule. The rabbit on the left comes from a cooler environment and is compact with short limbs and ears. The rabbit on the right comes from a warm environment and has long and lanky limbs and ears. The rabbit in the middle has ears and limbs that are in-between the other two. Rabbits do not sweat like humans; heat is dissipated primarily through their ears. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-13\/\">Allen\u2019s Rule (Figure 14.16b)<\/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<figure style=\"width: 627px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image17-6.png\" alt=\"Four noses with varying heights, arches, and nostril breadth.\" width=\"627\" height=\"237\" \/><figcaption class=\"wp-caption-text\">Figure 15.18: Human nasal morphological variation as influenced by four major climate-based adaptive zones: hot-dry, hot-wet, cold-dry, and cold-wet. The four noses in this figure vary in shape in relation to their respective climate-based adaptive zones. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-13\/\">Human nasal morphological variation (Figure 14.17)<\/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>Nasal shape and size (Figure 15.18) is another physiological feature affected by our ancestors\u2019 environments. The selective role of climate in determining human nasal variation is typically approached by dividing climates into four adaptive zones: hot-dry, hot-wet, cold-dry, and cold-wet (Maddux Et al. 2016). A principal role of the nasal cavity is to condition (i.e., warm and humidify) ambient air prior to its reaching the lungs. Given this function of the nasal cavity, it is anticipated that different nasal shapes and sizes will be related to varying environments. In cold-dry climates, an individual\u2019s nasal cavity must provide humidification and warmth to the dry air when breathing in through the nose (Noback Et al. 2011). Also, in that type of climate, the nasal cavity must conserve moisture and minimize heat loss during when the individual exhales through the nose (Noback Et al. 2011). From a physiological stress perspective, this is a stressful event.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Conversely, in hot-wet environments, there is no need for the nasal cavity to provide additional moisture to the inhaled air nor is there a need to warm the air or to preserve heat within the nasal cavity (Noback Et al. 2011). So, in hot-wet climates, the body is under less physiological stress related to the inhalation of ambient air than in cold-dry climates. As with most human morphological elements, the shape and size of the nasal cavity occurs along a cline. Due to the environmental stressors of cold-dry environments requiring the humidification and warming of air through the nasal cavity, individuals indigenous to such environments tend to have taller (longer) noses with a reduced nasal entrance (nostril opening) size (Noback Et al. 2011). This general shape is referred to as leptorrhine, and it allows for a larger surface area within the nasal cavity itself for the air to be warmed and humidified prior to entering the lungs (Maddux Et al. 2016). In addition, the relatively small nasal entrance of leptorrhine noses serves as a means of conserving moisture and heat (Noback Et al. 2011). Individuals indigenous to hot-wet climates tend to have platyrrhine nasal shapes, which are shorter with broader nasal entrances (Maddux Et al. 2016). Since individuals in hot-wet climates do not need to humidify and warm the air entering the nose, their nasal tract is shorter and the nasal entrance wider to permit the effective cooling of the nasal cavity during respiratory processes.<\/p>\n<h4 class=\"import-Normal\"><em>Adaptation: Infectious Disease<\/em><\/h4>\n<p class=\"import-Normal\">Throughout our evolutionary journey, humans have been exposed to numerous infectious diseases. In the following section, we will explore some of the evolutionary-based adaptations that have occurred in certain populations in response to the stressors presented by select infectious diseases. One of the primary examples of natural selection processes acting on the human genome in response to the presence of an infectious disease is the case of the relationship between the sickle-cell anemia trait and malaria, introduced in Chapter 5.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Malaria is a zoonotic disease (an infectious disease transmitted between animals and humans; it is covered in more detail in Chapter 17). It is caused by the spread of the parasitic protozoa from the genus <em>Plasmodium <\/em>(Figure 15.19). These unicellular, eukaryotic protozoa are transmitted through the bite of a female <em>Anopheles<\/em> mosquito. During the bite process, the protozoan parasites present within an infected mosquito\u2019s saliva enter a host\u2019s bloodstream where they are transported to the liver. Within the liver, the parasites multiply and are eventually released into the bloodstream, where they infect erythrocytes. Once inside the erythrocytes, the parasites reproduce until they exceed the cell\u2019s storage capacity, causing it to burst and release the parasites into the bloodstream once again. This replication cycle continues as long as there are viable erythrocytes within the host to infect.<\/p>\n<figure style=\"width: 518px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image13-1-1.jpg\" alt=\"Life cycle stages of Malaria parasite.\" width=\"518\" height=\"511\" \/><figcaption class=\"wp-caption-text\">Figure 15.19: Life cycle of the malaria parasite. <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:Malaria_parasite_life_cycle-NIAID.jpg\">Malaria parasite life cycle-NIAID<\/a> by <a href=\"https:\/\/www.niaid.nih.gov\/\">NIH National Institute of Allergy and Infectious Diseases<\/a> is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">General complications from malaria infections include the following: enlargement of the spleen (due to destruction of infected erythrocytes); lower number of thrombocytes (also called platelets, required for coagulation\/clotting of blood); high levels of bilirubin (a byproduct of hemoglobin breakdown in the liver) in the blood; jaundice (yellowing of the skin and eyes due to increased blood bilirubin levels); fever; vomiting; retinal (eye) damage; and convulsions (seizures). In 2020, there were 241 million cases of malaria reported globally, with 95% of those cases originating in Africa (World Health Organization 2021). In sub-Saharan Africa, where incidents of malaria are the highest in the world, 125 million pregnancies are affected by malaria, resulting in 200,000 infant deaths (Hartman, Rogerson, and Fischer 2013). Pregnant people who become infected during the gestational process are more likely to have low-birthweight infants due to prematurity or growth restriction inside the uterus (Hartman, Rogerson, and Fischer 2013). After birth, infants born to malaria-infected pregnant people are more likely to develop infantile anemia (low red-blood cell counts), a malaria infection that is not related to the maternal malarial infection, and they are more likely to die than infants born to non-malaria-infected pregnant people (Hartman, Rogerson, and Fischer 2013).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">For children and adolescents whose brains are still developing, there is a risk of cognitive (intellectual) impairment associated with some forms of malaria infections (Fernando, Rodrigo, and Rajapakse 2010). Given the relatively high rates of morbidity (disease) and mortality (number of deaths) associated with malaria, it is plausible that this disease may have served as a selective pressure during human evolution. Support for natural selection related to malaria resistance is related to genetic mutations associated with sickle cell, thalassemia, glucose-6-phosphate dehydrogenase (G6PD) deficiency, and the absence of certain antigens (molecules capable of inducing an immune response from the host) on erythrocytes. For the purposes of this text, we will focus our discussion on the relationship between sickle cell disease and malaria.<\/p>\n<figure style=\"width: 312px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image15-2-1.jpg\" alt=\"Normal (round) and sickle (crescent-shaped) red blood cells flow through a blood vessel. \" width=\"312\" height=\"499\" \/><figcaption class=\"wp-caption-text\">Figure 15.20: Normal and sickled erythrocytes. Normal red blood cells are round and can easily flow freely through blood vessels. Abnormal, or cicle, blood cells are half moon shaped and can easily become entangled and block blood flow. Sickle cells have abnormal hemoglobin that form strands that cause the sickle shape. The Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Sickle_cell_01.jpg\">Sickle cell 01<\/a> by <a href=\"https:\/\/www.nhlbi.nih.gov\/health\/sickle-cell-disease\">The National Heart, Lung, and Blood Institute (NHLBI)<\/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\"><strong>Sickle cell disease<\/strong> is a group of genetically inherited blood disorders characterized by an abnormality in the shape of the hemoglobin within erythrocytes. It is important to note that there are multiple variants of hemoglobin, including, but not limited to the following: A, D, C, E, F, H, S, Barts, Portland, Hope, Pisa, and Hopkins. Each of these variants of hemoglobin may result in various conditions within the body; however, for the following explanation we will focus solely on variants A and S.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Individuals who inherit a mutated gene (hemoglobin with a sickled erythrocyte variety, HbS) on chromosome 11 from both parents will develop sickle cell anemia, which is the most severe form of the sickle cell disease family (Figure 15.20). The genotype of an individual with sickle cell anemia is HbSS; whereas, an individual without sickle cell alleles has a genotype of HbAA representing two normal adult hemoglobin type A variants. Manifestations of sickle cell anemia (HbSS) range from mild to severe, with some of the more common symptoms being anemia, blood clots, organ failure, chest pain, fever, and low blood-oxygen levels. In high-income countries with advanced medical care, the median life expectancy of an HbSS individual is around 60 years; however, in low-income countries where advanced medical care is scarce, as many as 90% of children with sickle cell disease perish before the age of five (Longo Et al. 2017).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Considering that advanced medical care was not available during much of human evolutionary history, it stands to reason that the majority of individuals with the HbSS genotype died before the age of reproduction. If that is the case though, why do we still have the HbS variant present in modern populations? As covered earlier in this textbook, the genotype of an individual is composed of genes from both biological parents. In the case of an individual with an HbSS genotype, the sickle cell allele (HbS) was inherited from each of the parents. For individuals with the heterozygous genotype of HbSA, they have inherited both a sickle cell allele (HbS) and a normal hemoglobin allele (HbA). Heterozygous (HbSA) individuals who reside in regions where malaria is endemic may have a selective advantage. They will experience a sickling of some, but not all, of their erythrocytes. As discussed in the following paragraph, HbSA heterozygous individuals are less likely to die from malaria infections than their HbAA counterparts. Unlike an individual with the HbSS genotype, someone with HbSA may experience some of the symptoms listed above; however, they are generally less severe.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">As noted earlier, the mechanism through which <em>Plasmodium<\/em> protozoan parasites replicate involves human erythrocyte cells. However, due to their sickled shape, as well as the presence of an abnormally shaped protein within the cell, the parasites are unable to replicate effectively in the erythrocyte cells coded for by the HbS allele (Cyrklaff Et al. 2011). An individual who has an HbSA genotype and an active malaria infection will become ill with the disease to a lesser extent than someone with an HbAA genotype, which increases their chances of survival. Although normal erythrocytes (regulated by the HbA allele) allow for parasite replication, they are not able to replicate in HbS erythrocytes of the heterozygote. So, individuals with the HbSA genotype are more likely to survive a malaria infection than an individual who is HbAA. Although individuals with the HbSA genotype may endure some physiological complications related to the sickling of some of their erythrocytes, their morbidity and mortality rates are lower than they are for HbSS members of the population. The majority of individuals who are heterozygous or homozygous for the HbS trait have ancestors who originated in sub-Saharan Africa, India, Saudi Arabia, and regions in South and Central America, the Mediterranean (Turkey, Greece, and Italy), and the Caribbean (Centers for Disease Control and Prevention 2017; Figure 15.21).<\/p>\n<figure style=\"width: 511px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image23-3.png\" alt=\"Map illustrating distribution of sickle cell and associated erythrocytic abnormalities for Africa and Asia.\" width=\"511\" height=\"306\" \/><figcaption class=\"wp-caption-text\">Figure 15.21: Distribution of sickle cell and associated erythrocytic abnormalities for Africa and Asia. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Red_Blood_Cell_abnormalities.png\">Red Blood Cell abnormalities<\/a> by Armando Moreno Vranich 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\">With respect to the history of these regions, during the early phases of settlement horticulture was the primary method of crop cultivation. Typically performed on a small scale, horticulture is based on manual labour and relatively simple hand tools rather than the use of draft animals or irrigation technologies. Common in horticulture is <em>swidden<\/em>, or the cutting and burning of plants in woodland and grassland regions. The swidden is the prepared field that results following a slash-and-burn episode. This practice fundamentally alters the soil chemistry, removes plants that provide shade, and increases the areas where water may pool. This anthropogenically altered landscape provides the perfect breeding ground for the <em>Anopheles<\/em> mosquito, as it prefers warm, stagnant pools of water (Figure 15.22).<\/p>\n<figure style=\"width: 561px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5-6.png\" alt=\"Horticulture encourages mosquitos, increasing malaria. Balancing selection maintains normal and sickle cell alleles.\" width=\"561\" height=\"468\" \/><figcaption class=\"wp-caption-text\">Figure 15.22: The effects of human horticultural activities on the balancing selection of populations in relation to sickle cell disease genotype variants. <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__-13\/\">Sickle cell disease (Figure 14.21)<\/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>. [Includes two horticulture illustrations by Mary Nelson, <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Sickle_cell_anemia.jpg\">Sickle cell anemia<\/a> by <a href=\"https:\/\/en.wikibooks.org\/wiki\/User:Pkleong\">Pkleong<\/a> at <a href=\"https:\/\/en.wikibooks.org\/wiki\/\">English Wikibooks<\/a> (modified), <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\">Although swidden agriculture was historically practiced across the globe, it became most problematic in the regions where the <em>Anopheles<\/em> mosquito is endemic. These areas have the highest incidence rates of malaria infection. Over time, the presence of the <em>Anopheles<\/em> mosquito and the <em>Plasmodium<\/em> parasite that it transmitted acted as a selective pressure, particularly in regions where swidden agricultural practices were common, toward the selection of individuals with some modicum of resistance against the infection. In these regions, HbSS and HbSA individuals would have been more likely to survive and reproduce successfully. Although individuals and populations are far more mobile now than they have been throughout much of history, there are still regions where we can see higher rates of malaria infection as well as greater numbers of individuals with the HbS erythrocyte variant. The relationship between malaria and the selective pressure for the HbS variant is one of the most prominent examples of natural selection in the human species within recent evolutionary history.<\/p>\n<h4 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><em>Adaptation: Lactase Persistence <\/em><\/h4>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">With the case of sickled erythrocytes and their resistance to infection by malaria parasites, there is strong support for a cause-and-effect-style relationship linked to natural selection. Although somewhat less apparent, there is a correlation between lactase persistence and environmental challenges. Lactase-phlorizin hydrolase (LPH) is an enzyme that is primarily produced in the small intestine and permits the proper digestion of lactose, a disaccharide (composed of two simple sugars: glucose and galactose) found in the milk of mammals. Most humans will experience a decrease in the expression of LPH following weaning, leading to an inability to properly digest lactose. Generally, LPH production decreases between the ages of two and five and is completely absent by the age of nine (Dzialanski Et al. 2016). For these individuals, the ingestion of lactose may lead to a wide variety of gastrointestinal ailments, including abdominal bloating, increased gas, and diarrhea. Although the bloating and gas are unpleasant, the diarrhea caused by a failure to properly digest lactose can be life-threatening if severe enough due to the dehydration it can cause. Some humans, however, are able to produce LPH far beyond the weaning period.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Individuals who continue to produce LPH have what is referred to as the <strong>lactase persistence <\/strong>trait. The lactase persistence trait is encoded for a gene called <em>LCT<\/em>, which is located on human chromosome 2 (Ranciaro Et al. 2014). From an evolutionary and historical perspective, this trait is most commonly linked to cultures that have practiced cattle domestication (Figure 15.23). For individuals in those cultures, the continued expression of LPH may have provided a selective advantage. During periods of environmental stress, such as a drought, if an individual is capable of successfully digesting cow\u2019s milk, they have a higher chance of survival than someone who suffers from diarrhea-linked dehydration due to a lack of LPH. Although the frequency of the lactase persistence trait is relatively low among African agriculturalists, it is high among pastoralist populations that are traditionally associated with cattle domestication, such as the Tutsi and Fulani, who have frequencies of 90% and 50%, respectively (Tishkoff Et al. 2007).<\/p>\n<figure style=\"width: 632px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-4.png\" alt=\"Map depicting the percentage of adults with lactase persistence in the eastern hemisphere.\" width=\"632\" height=\"497\" \/><figcaption class=\"wp-caption-text\">Figure 15.23: Interpolated map depicting the percentage of adults with the lactase persistence genotype in indigenous populations of Africa, Europe, Asia, and Australia. Circles denote sample locations. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Lactose_tolerance_in_the_Old_World.svg\">Lactose tolerance in the Old World<\/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\/4.0\/legalcode\">CC BY 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Cattle domestication began around 11,000 years ago in Europe (Beja-Pereira Et al. 2006) and 7,500 to 9,000 years ago in the Middle East and North Africa (Tishkoff Et al. 2007). Based on human genomic studies, it is estimated that the mutation for the lactase persistence trait occurred around 2,000 to 20,000 years ago for European populations (Tishkoff Et al. 2007). For African populations, the lactase persistence trait emerged approximately 1,200 to 23,000 years ago (Gerbault Et al. 2011). This begs the question: Is this mutation the same for both populations? It appears that the emergence of the lactase persistence mutation in non-European populations, specifically those in East Africa (e.g., Tutsi and Fulani), is a case of <strong>convergent evolution<\/strong>. With convergent evolution events, a similar mutation may occur in species of different lineages through independent evolutionary processes. Based on our current understanding of the genetic mutation pathways for the lactase persistence trait in European and African populations, these mutations are not representative of a shared lineage. In other words, just because a person of European origin and a person of African origin can each digest milk due to the presence of the lactase-persistence trait in their genotypes, it does not mean that these two individuals inherited it due to shared common ancestry.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Is it possible that the convergent evolution of similar lactase-persistence traits in disparate populations is merely a product of genetic drift? Or is there evidence for natural selection? Even though 23,000 years may seem like a long time, it is but a blink of the proverbial evolutionary eye. From the perspective of human evolutionary pathways, mutations related to the <em>LCT<\/em> gene have occurred relatively recently. Similar genetic changes in multiple populations through genetic drift processes, which are relatively slow and directionless, fail to accumulate as rapidly as lactase-persistence traits (Gerbault Et al. 2011). The widespread accumulation of these traits in a relatively short period of time supports the notion that an underlying selective pressure must be driving this form of human evolution. Although to date no definitive factors have been firmly identified, it is thought that environmental pressures are likely to credit for the rapid accumulation of the lactase-persistence trait in multiple human populations through convergent evolutionary pathways.<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Skin Tone Genetic Regulation<\/h2>\n<p class=\"import-Normal\">The melanocortin 1 receptor (<em>MC1R<\/em>) gene acts to control which types of melanin (eumelanin or pheomelanin) are produced by melanocytes. The <em>MC1R<\/em> receptor is located on the surface of the melanocyte cells (Quillen Et al. 2018). Activation of the <em>MC1R <\/em>receptors may occur through exposure to specific environmental stimuli or due to underlying genetic processes. Inactive or blocked <em>MC1R <\/em>receptors result in melanocytes producing pheomelanin. If the <em>MC1R<\/em> gene receptors are activated, then the melanocytes will produce eumelanin. Thus, individuals with activated <em>MC1R<\/em> receptors tend to have darker-pigmented skin and hair than individuals with inactive or blocked receptors.<\/p>\n<p class=\"import-Normal\">The alleles of another gene, the major facilitator, superfamily domain-containing protein 12 (<em>MFSD12<\/em>) gene, affect the expression of melanocytes in a different way than the <em>MC1R<\/em> gene. Instead of affecting the activation of melanocyte receptors, the <em>MFSD12<\/em> alleles indirectly affect the membranes of melanocyte lysosomes (Quillen Et al. 2018). The melanocyte\u2019s lysosomes are organelles containing digestive enzymes, which ultimately correlate to varying degrees of pigmentation in humans. Variations in the membranes of the melanocyte lysosomes ultimately correlate to differing degrees of pigmentation in humans.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ancestral <em>MFSD12 <\/em>allele variants are present in European and East Asian populations and are associated with lighter pigmentation of the skin (Crawford Et al. 2017; Quillen Et al. 2018). In addition, this ancestral variant is also associated with Tanzanian, San, and Ethiopian populations of Afro-Asiatic ancestry (Crawford Et al. 2017; Quillen Et al. 2018). In contrast, the more derived (i.e., more recent) allele variants that are linked to darker skin tones are more commonly present in East African populations, particularly those of Nilo-Saharan descent (Crawford Et al. 2017; Quillen Et al. 2018). The notion that ancestral alleles of <em>MFSD12<\/em> are associated with lighter skin pigmentation is in opposition to the commonly accepted idea that our pigmentation was likely darker throughout early human evolution (Crawford Et al. 2017; Quillen Et al. 2018). Due to the complexity of the human genome, <em>MFSD12<\/em> and <em>MC1R <\/em>are but two examples of alleles affecting human skin tone. Furthermore, there is genetic evidence suggesting that certain genomic variants associated with both darker and lighter skin colour have been subject to directional selection processes for as long as 600,000 years, which far exceeds the evolutionary span of <em>Homo sapiens sapiens<\/em> (Crawford Et al. 2017; Quillen Et al. 2018).<\/p>\n<\/div>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<div class=\"textbox shaded\">\n<h2>Summary<\/h2>\n<p>From the time that the first of our species left Africa, we have had to adjust and adapt to numerous environmental challenges. The remarkable ability of human beings to maintain homeostasis through a combination of both nongenetic (adjustments) and genetic (adaptations) means has allowed us to occupy a remarkable variety of environments, from high-altitude mountainous regions to the tropics near the equator. From adding piquant, pungent spices to our foods as a means of inhibiting food-borne illnesses due to bacterial growth to donning garments specially suited to local climates, behavioural adjustments have provided us with a nongenetic means of coping with obstacles to our health and well-being. Acclimatory adjustments, such as sweating when we are warm in an attempt to regulate our body temperature or experiencing increased breathing rates as a means of increasing blood oxygen levels in regions where the partial pressure of oxygen is low, have been instrumental in our survival with respect to thermal and altitudinal environmental challenges. For some individuals, developmental adjustments that were acquired during their development and growth phases (e.g., increased heart and lung capacities for individuals from high-altitude regions) provide them with a form of physiological advantage not possible for someone who ventures to such an environmentally challenging region as an adult. Genetically mediated adaptations, such as variations in the pigmentation of our skin, have ensured our evolutionary fitness across all latitudes. Taken together, these examples point to a broader question: will the human species continue to adjust and adapt to new environmental challenges in the future? If past performance is any measure of future expectations, then the human story will continue as long as we do not alter our environment to the point that the plasticity of our behaviour, physiological, and morphological boundaries is exceeded.<\/p>\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\">Detail at least two examples of how natural selection has influenced human variation. Specifically, what was the selective pressure that may have led to a preference for a specific trait and how is that trait related to an increased level of fitness?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Why is reduced pigmentation of the skin advantageous for individuals from northern latitudes? What role does darker skin pigmentation serve for individuals near the equator? What is the relationship between skin pigmentation and fitness?<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">What are some of the risks associated with pregnancy at high altitude? Compare and contrast the various genetic mutations of the indigenous Tibetan and Ethiopian high-altitude populations. In your answer, specifically address the issue of pregnancy at high altitudes.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">What is the relationship between the sickle cell mutation and the <em>Plasmodium<\/em> parasite? Would having the HbSA genotype still be advantageous in a region where such parasites are not common? Why or why not?<\/li>\n<\/ul>\n<\/div>\n<p>&nbsp;<\/p>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Key Terms<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Acclimatory adjustm<\/strong><strong>ents<\/strong>: Processes by which an individual organism adjusts in order to maintain homeostasis in response to environmental challenges.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Activated melanogenesis<\/strong>: Increase in melanin production in response to ultraviolet radiation (UV) exposure.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Adaptation<\/strong>: Alteration in population-level gene frequencies related to environmentally induced selective pressures; leads to a greater level of fitness for a population related to a specific environment.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Adjustment<\/strong>: Nongenetic-based ways in which organisms adjust to environmental stressors.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Allen\u2019s Rule<\/strong>: Due to thermal adaptation, homeothermic animals have body volume-to-surface ratios that vary inversely with the average temperature of their environment. In cold climates, the anticipated ratio is high; in warm climates, it is low.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Basal melanogenesis<\/strong>: Genetically mediated, non-environmentally influenced base melanin level.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Behavioural adjustments<\/strong>: An individual\u2019s culturally mediated responses to an environmental stressor in an effort to maintain homeostasis.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Bergmann\u2019s Rule<\/strong>: For a broadly distributed monophyletic group, species and populations of smaller size tend to be found in environments with warmer climates and those of larger size tend to be found in ones that are colder.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Cline<\/strong>: A continuum of gradations (i.e., degrees or levels) of a specific trait.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Conduction<\/strong>: Mechanism of heat transfer between objects through direct contact.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Convection<\/strong>: Movement of heat away from a warm object to the cooler surrounding fluid (i.e., gas or liquid).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Convergent evolution<\/strong>: Evolutionary process whereby organisms that are not closely related independently evolve similar traits as a product of adaptation to similar evolutionary parameters.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Erythrocyte<\/strong>: Red blood cell; most common form of blood cell; the principle means of transporting oxygen throughout the circulatory system.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Evaporation<\/strong>: Mechanism of heat transfer whereby liquid is transformed into a gas, utilizing energy (e.g., heat).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Folic acid<\/strong>: Form of B complex vitamin necessary for proper fetal development.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Gloger\u2019s Rule<\/strong>: For mammals of the same species, those with more darkly pigmented forms tend to be found closer to the equator and those with lighter forms are found in regions further from the equator.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Homeostasis<\/strong>: Condition of optimal functioning for an organism.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Hyperpnea<\/strong>: Increased depth and rate of respiration.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Hypothalamus<\/strong>: Small portion of the human brain responsible for body temperature regulation.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Lactase persistence<\/strong>: Genetic mutation permitting the continued production of lactase-phlorizin hydrolase enzyme in the small intestine past the weaning period.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Melanin<\/strong>: Black-brown pigment produced by melanocytes; one of the primary pigments in skin.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Melanocytes<\/strong>: Specialized cells that produce melanin.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Phenotypic plasticity<\/strong>: Ability of one genotype to produce more than one phenotype dependent on environmental conditions.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Radiation<\/strong>: Mechanism of heat transfer involving electromagnetic energy being emitted from an object.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Sickle cell disease<\/strong>: A group of genetically inherited blood disorders characterized by an abnormality in the shape of the hemoglobin within erythrocytes (red blood cells).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Stressor<\/strong>: Any stimulus resulting in an imbalance in an organism\u2019s homeostatic balance.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Vasoconstriction<\/strong>: Narrowing of the blood vessels due to contractions of the muscular vessel walls.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Vasodilation<\/strong>: Dilation of the blood vessels due to relaxation of the muscular vessel walls.<\/p>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">For Further Exploration<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Homeostasis<\/strong><\/p>\n<p class=\"import-Normal\">Baptista, Vander. 2006. \u201c<a href=\"https:\/\/doi.org\/10.1152\/advan.00075.2006\" target=\"_blank\" rel=\"noopener\">Starting Physiology: Understanding Homeostasis<\/a>.\u201d <em>Advances in Physiology Education<\/em> 30: 263\u2013264.<\/p>\n<p class=\"import-Normal\">Goldstein, David S., and Bruce McEwen. 2002. \u201c<a href=\"https:\/\/doi.org\/10.1080\/102538902900012345\" target=\"_blank\" rel=\"noopener\">Allostasis, Homeostats, and the Nature of Stress<\/a>.\u201d <em>The International Journal on the Biology of Stress<\/em> 5 (1): 55\u201358.<\/p>\n<p class=\"import-Normal\"><strong>General Clinal Variation and Genetic Exchange<\/strong><\/p>\n<p class=\"import-Normal\">Delhey, Kaspar. 2019. \u201c<a href=\"https:\/\/doi.org\/10.1111\/brv.12503\" target=\"_blank\" rel=\"noopener\">A Review of Gloger's Rule, an Ecogeographical Rule of Colour: Definitions, Interpretations and Evidence<\/a>.\u201d <em>Biological Reviews<\/em> 94 (4): 1294\u20131316.<\/p>\n<p class=\"import-Normal\">Feng, Yuanqing, Michael A. McQuillan, and Sarah A. Tishkoff. 2021. \u201c<a href=\"https:\/\/doi.org\/10.1093\/hmg\/ddab007\" target=\"_blank\" rel=\"noopener\">Evolutionary Genetics of Skin Pigmentation in African Populations<\/a>.\u201d <em>Human Molecular Genetics<\/em> 30 (R1): R88\u2013R97.<\/p>\n<p class=\"import-Normal\">Hu, Hao, Nayia Petousi, Gustavo Glusman, Yao Yu, Ryan Bohlender, Tsewang Tashi, Jonathan M. Downie, et al. 2017. \u201c<a href=\"https:\/\/doi.org\/10.1371\/journal.pgen.1006675\" target=\"_blank\" rel=\"noopener\">Evolutionary History of Tibetans Inferred from Whole-Genome Sequencing<\/a>.\u201d <em>PLoS Genetics <\/em>13 (4): e1006675. .<\/p>\n<p class=\"import-Normal\">Jablonski, Nina G. 2021. \u201c<a href=\"https:\/\/doi.org\/10.1002\/ajpa.24200\" target=\"_blank\" rel=\"noopener\">Skin Color and Race<\/a>.\u201d Special issue, \u201cRace Reconciled II: Interpreting and Communicating Biological Variation and Race in 2021,\u201d <em>American Journal of Physical Anthropology<\/em> 175 (2): 437\u2013447.<\/p>\n<p class=\"import-Normal\">Pritchard, Jonathan K., Joseph K. Pickrell, and Graham Coop. 2010. \u201c<a href=\"https:\/\/doi.org\/10.1016\/j.cub.2009.11.055\" target=\"_blank\" rel=\"noopener\">The Genetics of Human Adaptation: Hard Sweeps, Soft Sweeps, and Polygenic Adaptation<\/a>.\u201d <em>Current Biology<\/em> 20 (4): R208\u2013R215.<\/p>\n<p class=\"import-Normal\">Sankararaman, Sriram, Swapan Mallick, Nick Patterson, and David Reich. 2016. \u201c<a href=\"https:\/\/doi.org\/10.1016\/j.cub.2016.03.037\" target=\"_blank\" rel=\"noopener\">The Combined Landscape of Denisovan and Neanderthal Ancestry in Present-Day Humans<\/a>.\u201d <em>Current Biology<\/em> 26 (9): 1241\u20131247.<\/p>\n<p class=\"import-Normal\"><strong>Lactase Persistence<\/strong><\/p>\n<p class=\"import-Normal\">HHMI BioInteractive. 2021. \u201c<a href=\"https:\/\/www.biointeractive.org\/classroom-resources\/making-fittest-got-lactase-co-evolution-genes-and-culture\" target=\"_blank\" rel=\"noopener\">The Making of the Fittest: Got Lactase? The Co-evolution of Genes and Culture<\/a>.\u201d Accessed April 7, 2023.<\/p>\n<p class=\"import-Normal\"><strong>Malaria and Sickle Cell Anemia<\/strong><\/p>\n<p class=\"import-Normal\">Bill and Melinda Gates Foundation. 2022. \u201c<a href=\"https:\/\/www.gatesfoundation.org\/our-work\/programs\/global-health\/malaria\" target=\"_blank\" rel=\"noopener\">Malaria<\/a>.\u201d Accessed April 7, 2023.<\/p>\n<p class=\"import-Normal\">Centers for Disease Control and Prevention. 2022. \u201c<a href=\"https:\/\/www.cdc.gov\/parasites\/malaria\/index.html\" target=\"_blank\" rel=\"noopener\">Malaria<\/a>.\u201d Accessed April 7, 2023.<\/p>\n<p class=\"import-Normal\">HHMI BioInteractive. 2020. \u201c<a href=\"https:\/\/www.biointeractive.org\/classroom-resources\/making-fittest-natural-selection-humans\" target=\"_blank\" rel=\"noopener\">The Making of the Fittest: Natural Selection in Humans<\/a>.\u201d 2020. Accessed April 7, 2023.<\/p>\n<p class=\"import-Normal\">National Institutes of Health: National Center for Advancing Translational Sciences. \u201c<a href=\"https:\/\/rarediseases.info.nih.gov\/diseases\/8614\/sickle-cell-anemia\" target=\"_blank\" rel=\"noopener\">Sickle Cell Anemia<\/a>.\u201d Accessed April 7, 2023.<\/p>\n<p class=\"import-Normal\">World Health Organization. 2022. \u201c<a href=\"https:\/\/www.who.int\/news-room\/fact-sheets\/detail\/malaria\" target=\"_blank\" rel=\"noopener\">Malaria<\/a>.\u201d Accessed April 7, 2023.<\/p>\n<p class=\"import-Normal\"><strong>Rickets and Bone Health<\/strong><\/p>\n<p class=\"import-Normal\">National Institutes of Health: National Center for Advancing Translational Sciences. \u201c<a href=\"https:\/\/rarediseases.info.nih.gov\/diseases\/5700\/rickets\" target=\"_blank\" rel=\"noopener\">Rickets<\/a>.\u201d Accessed April 7, 2023.<\/p>\n<p class=\"import-Normal\">Talmadge, D. W., and R. V. Talmadge. 2007. \u201cCalcium Homeostasis: How Bone Solubility Relates to All Aspects of Bone Physiology.\u201d <em>Journal of Musculoskeletal and Neuronal Interactions <\/em>7 (2): 108\u2013112.<\/p>\n<p class=\"import-Normal\"><strong>Skin Colour<\/strong><\/p>\n<p class=\"import-Normal\">HHMI BioInteractive. 2020. \u201c<a href=\"https:\/\/www.biointeractive.org\/classroom-resources\/biology-skin-color\" target=\"_blank\" rel=\"noopener\">The Biology of Skin Color<\/a>.\u201d Accessed April 7, 2023<\/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\">American Academy of Pediatrics, Task Force on Infant Sleep Position and Sudden Infant Death Syndrome. 2000. \u201cChanging Concepts of Sudden Infant Death Syndrome: Implications for Infant Sleeping Environment and Sleep Position.\u201d <em>Pediatrics<\/em> 105 (3): 650\u2013656.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Beja-Pereira, Albano, David Caramelli, Carles Lalueza-Fox, Cristiano Vernesi, Nuno Ferrand, Antonella Casoli, Felix Goyache, et al. 2006. \u201cThe Origin of European Cattle: Evidence from Modern and Ancient DNA.\u201d <em>PNAS <\/em>103 (21): 8113\u20138118.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Best, Andre, Daniel E. Lieberman, and Jason M. Kamilar. 2019. \u201cDiversity and Evolution of Human Eccrine Sweat Gland Density.\u201d <em>Journal of Thermal Biology<\/em> 84: 331\u2013338.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Bigham, Abigail W. 2016. \u201c<a href=\"https:\/\/www.cdc.gov\/ncbddd\/sicklecell\/data.html\" target=\"_blank\" rel=\"noopener\">Genetics of Human Origin and Evolution: High-Altitude Adaptations<\/a>.\u201d <em>Current Opinion in Genetics &amp; Development<\/em> 41: 8\u201313.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Centers for Disease Control and Prevention. 2017. \u201cData &amp; Statistics on Sickle Cell Disease.\u201d <em>Centers for Disease Control and Prevention<\/em> website, August 9. Accessed April 7, 2023. .<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Crawford, Nicholas G., Derek E. Kelly, Matthew E. B. Hansen, Marcia H. Beltrame, Shaohua Fan, Shanna L. Bowman, Ethan Jewett, et al. 2017. \u201cLoci Associated with Skin Pigmentation Identified in African Populations.\u201d <em>Science<\/em> 358 (6365): 1\u201349.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Cyrkalff, Marek, Cecilia P. Sanchez, Nicole Kilian, Curille Bisseye, Jacques Simpore, Friedrich Frischknecht, and Michael Lanzer. 2011. \u201cHemoglobins S and C Interfere with Actin Remodeling in <em>Plasmodium <\/em><em>falciparum<\/em>-Infected Erythrocytes.\u201d <em>Science<\/em> 334 (6060): 1283\u20131286.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Dzialanski, Zbigniew, Michael Barany, Peter Engfeldt, Anders Magnuson, Lovisa A. Olsson, and Torbj\u04e7rn K. Nilsson. 2016. \u201cLactase Persistence versus Lactose Intolerance: Is There an Intermediate Phenotype?\u201d <em>Clinical Biochemistry<\/em> 49 (2016): 248\u2013252.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Fernando, Sumadya D., Chaturaka Rodrigo, and Senaka Rajapakse. 2010. \u201cThe \u2018Hidden\u2019 Burden of Malaria: Cognitive Impairment Following Infection.\u201d <em>Malaria Journal<\/em> 9 (366): 1\u201311.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Gerbault, Pascale, Anke Liebert, Yuval Itan, Adam Powell, Mathias Currat, Joachim Burger, Dallas M. Swallow, and Mark G. Thomas. 2011. \u201cEvolution of Lactase Persistence: An Example of Human Niche Construction.\u201d <em>Philosophical Transactions of the Royal Society B: Biological Sciences <\/em>366 (1566): 863\u2013877.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Gray, Olivia A., Jennifer Yoo, D\u00e9bora R. Sobriera, Jordan Jousma, David Witnosky, Noboru J. Sakabe, Ying-Jie Ping, et al. 2022. \u201cA Pleiotropic Hypoxia-Sensitive <em>EPAS1<\/em> Enhancer Is Disrupted by Adaptive Alleles in Tibetans.\u201d <em>Science Advances <\/em>8 (47): 1\u201313.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Hartman, T. K., S. J. Rogerson, and P. R. Fischer. 2013. \u201cThe Impact of Maternal Malaria on Newborns.\u201d <em>Annals of Tropical Paediatrics<\/em> 30 (4): 271\u2013282.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Longo, Dan L., Fr\u00e9d\u00e9ric B. Piel, Martin H. Steinberg, and David C. Rees. 2017. \u201cSickle Cell Disease.\u201d <em>The New England Journal of Medicine<\/em> 376 (16): 1561\u20131573.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Maddux, Scott D., Todd R. Yokley, Bohumil M. Svoma, and Robert G. Franciscus. 2016. \u201cAbsolute Humidity and the Human Nose: A Reanalysis of Climate Zones and Their Influence on Nasal Form and Function.\u201d <em>American Journal of Physical Anthropology <\/em>161 (2): 309\u2013320.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Meyer, M. C., M. S. Alexander, Z. Wang, D. L. Hoffmann, J. A. Dahl, D. Degering, W. R. Haas, and F. Schl\u00fctz. 2017. \u201cPermanent Human Occupation of the Central Tibetan Plateau in the Early Holocene.\u201d <em>Science <\/em>355 (6320): 64\u201367.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Moore, Lorna G., Susan Niermeyer, and Stacy Zamudio. 1998. \u201cHuman Adaptation to High Altitude: Regional and Life-Cycle Perspectives.\u201d <em>Yearbook of Physical Anthropology<\/em> 41: 25\u201364.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Noback, Marlijn L., Katerina Harvati, and Fred Spoor. 2011. \u201cClimate-Related Variation of the Human Nasal Cavity.\u201d <em>American Journal of Physical Anthropology<\/em> 145 (4): 599\u2013614.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Peacock, A. J. 1998. \u201cABC of Oxygen: Oxygen at High Altitude.\u201d <em>BMJ<\/em> 317 (7165): 1063\u20131066.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Pontzer, Herman, Mary H. Brown, Brian M. Wood, David A. Raichlen, Audax Z.P. Madbulla, Jacob A. Harris, Holly Dunsworth, et al. 2021. \u201cEvolution of Water Conservation in Humans.\u201d <em>Current Biology<\/em> 31 (8): 1804\u20131810.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Quillen, Ellen E., Heather L. Norton, Esteban J. Parra, Frida Loza-Durazo, Khai C. Ang, Florin Mircea Illiescu, Laurel N. Pearson, et al. 2019. \u201cShades of Complexity: New Perspectives on the Evolution and Genetic Architecture of Human Skin.\u201d <em>Yearbook<\/em><em> of Physical Anthropology <\/em>168 (S67): 4\u201326.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ranciaro, Alessia, Michael C. Campbell, Jibril B. Hirbo, Wen-Ya Ko, Alain Froment,<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Paolo Anagnostou, Maritha J. Kotze, et al. 2014. \u201cGenetic Origins of Lactase Persistence and the Spread of Pastoralism in Africa.\u201d <em>American Journal of Human Genetics <\/em>94 (4): 496\u2013510.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Roby, Brianne Barnett, Marsha Finkelstein, Robert J. Tibesar, and James D. Sidman. 2012. \u201cPrevalence of Positional Plagiocephaly in Teens Born after the \u2018Back to Sleep\u2019 Campaign.\u201d <em>Otolaryn<\/em><em>g<\/em><em>ology<\/em><em>\u2014Head and Neck Surgery<\/em> 146 (5): 823\u2013828.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Sherman, Paul W., and Jennifer Billing. 1999. \u201cDarwinian Gastronomy: Why We Use Spices.\u201d <em>BioScience <\/em>49 (6): 453\u2013463.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Tishkoff, Sarah A., Floyd A. Reed, Alessia Ranciaro, Benjamin F. Voight, Courtney C. Babbitt, Jesse S. Silverman, Kweli Powell, et al. 2007. \u201cConvergent Adaptation of Human Lactase Persistence in Africa and Europe.\u201d <em>Nature Genetics<\/em> 39 (1): 31\u201340.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">World Health Organization. 2021. \u201c<a href=\"https:\/\/www.who.int\/teams\/global-malaria-programme\/reports\/world-malaria-report-2021\" target=\"_blank\" rel=\"noopener\">World Malaria Report 2021<\/a>.\u201d <em>World Health Organization<\/em> website, December 4, 2022. Accessed April 7, 2023.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Zhang, Xinjun, Kelsey E. Witt, Mayra M. Ba\u00f1uelos, Amy Ko, Kai Yuan, Shuhua Xu, Rasmus Nielsen, and Emilia Huerta-Sanchez. 2021. \u201cThe History and Evolution of Denisovan-<em>EPAS1<\/em> Haplotype in Tibetans.\u201d <em>PNAS Biological Sciences<\/em> 118 (22): 1\u20139.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">\n<\/div>\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_1683_1694\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1694\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\">Michael B. C. Rivera, Ph.D., University of Cambridge<\/p>\n<p class=\"import-Normal\"><em>This chapter is a revision from <\/em><em>\"<\/em><a class=\"rId7\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-12\/\"><em>Chapter 13: Race and Human Variation<\/em><\/a><em>\u201d by Michael B. C. Rivera. 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\">Learning Objectives<\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li>Illustrate the troubling history of \u201crace\u201d concepts.<\/li>\n<li>Explain human variation and evolution as the thematic roots of biological anthropology as a discipline.<\/li>\n<li>Critique earlier \u201crace\u201d concepts based on a contemporary understanding of the apportionment of human genetic variation.<\/li>\n<li>Explain how biological variation in humans is distributed clinally and in accordance with both isolation-by-distance and Out-of-Africa models.<\/li>\n<li>Identify phenotypic traits that reflect selective and neutral evolution.<\/li>\n<li>Extend this more-nuanced view of human variation to today\u2019s research, the implications for biomedical studies, applications in forensic anthropology, and other social\/cultural\/political concerns.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p>Humans exhibit biological variation. Humans also have a universal desire to categorize other humans in order to make sense of the world around them. Since the birth of the discipline of <strong>biological anthropology, <\/strong>we have been interested in studying how humans vary biologically and what the sources of this variation are. Before we tackle these big problems, first consider this question: Why <em>should<\/em> we study human variation?<\/p>\n<figure style=\"width: 429px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2023\/06\/image26-3.png\" alt=\"Culturally and biologically diverse humans.\" width=\"429\" height=\"429\" \/><figcaption class=\"wp-caption-text\">Figure 14.1: Humans are culturally diverse (in that cultural differences contribute to a great degree of variation between individuals), but those shown are genetically undiverse. (Top left: Hadzabe members in Tanzania; top right: Inuit family; bottom left: Andean man in Peru; bottom right: English woman.) Credit: <a class=\"rId11\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-12\/\">Humans are diverse (Figure 13.1)<\/a> original to <a class=\"rId12\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Michael Rivera is a collective work under a <a class=\"rId13\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0<\/a> license. [Includes <a class=\"rId14\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Tanzania_-_Hadzabe_hunter_(14533536392).jpg\">Tanzania - Hadzabe hunter (14533536392)<\/a> by <a class=\"rId15\" href=\"https:\/\/www.flickr.com\/people\/67947877@N06\">A_Peach<\/a>, <a class=\"rId16\" href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\">CC BY 2.0<\/a>; <a class=\"rId17\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Inuit-Kleidung_1.jpg\">Inuit-Kleidung 1<\/a> by Ansgar Walk, <a class=\"rId18\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0<\/a>; <a class=\"rId19\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Andean_Man.jpg\">Andean Man<\/a> by <a class=\"rId20\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Cacophony\">Cacophony<\/a>, <a class=\"rId21\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0<\/a>; <a class=\"rId22\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Jane_Goodall_GM.JPG\">Jane Goodall GM<\/a> by <a class=\"rId23\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Floatjon\">Floatjon<\/a>, <a class=\"rId24\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">There are certainly academic reasons for studying human variation. First, it is highly interesting and important to consider the evolution of our species (see Chapters 10-13) and how our biological variation may be similar to (or different from) that of other species (e.g., other primates and apes; see Chapters 6 and 7). Second, anthropologists study modern human variation to understand how different biological traits developed over evolutionary time (see Figure 14.1). Suppose we are able to grasp the evolutionary processes that produce the differences in biology, physiology, body chemistry, behaviour, and culture (<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_804\">human variation<\/a><\/strong>). In that case, we can make more accurate inferences about evolution and adaptation among our hominin ancestors, complementing our study of fossil evidence and the archaeological record. Third, as will be discussed in more detail later on, it is important to consider that biological variation among humans has biomedical, forensic, and sociopolitical implications. For these reasons, the study of human variation and evolution has formed the basis of anthropological inquiry for centuries and continues to be a major source of intrigue and inspiration for scientific research conducted today.<\/p>\n<p class=\"import-Normal\">An even-more-important role of the biological anthropologist is to improve public understanding of human evolution and variation\u2014outside of academic circles. Terms such as <strong>race<\/strong> and <strong>ethnicity<\/strong> are used in everyday conversations and in formal settings within and outside academia. The division of humankind into smaller, discrete categories is a regular occurrence in day-to-day life. This can be seen regularly when governments acquire census data with a heading like \u201cgeographic origin\u201d or \u201cethnicity.\u201d Furthermore, such checkboxes and drop-down lists are commonly seen as part of the identifying information required for surveys and job applications.<\/p>\n<p class=\"import-Normal\">According to professors of anthropology, ethnic studies, and sociology, race is often understood as rooted in biological differences, ranging from such familiar traits as skin colour or eye shape to variations at the genetic level. However, race can also be studied as an \u201cideological construct\u201d that goes beyond biological and genetic bases (Fuentes Et al. 2019), at different times relating to our ethnicities, languages, religious beliefs, and cultural practices. Sometimes people associate racial identity with the concept of socioeconomic status or position, or they link ideas about race to what passport someone has, how long they have been in a country, or how well they have \u201cintegrated\u201d into a population.<\/p>\n<p class=\"import-Normal\">Some of these ideas about ethnicity have huge social and political impacts, and notions of race have been part of the motivation behind various forms of racism and prejudice today, as well as many wars and genocides throughout history. <strong>Racism<\/strong> manifests in many ways\u2014from instances of bullying between kids on school playgrounds to underpaid minorities in the workforce, and from verbal abuse hurled at people of colour to violent physical behaviours against those of a certain race. <strong>Prejudice<\/strong> can be characterized as negative views toward another group based on some perceived characteristic that makes all members of that group worthy of disdain, disrespect, or exclusion (not solely along racial lines but also according to [dis]ability, gender, sexual orientation, or socioeconomic background, for example). According to Shay-Akil McLean (2014), \u201cRacism is not something particular to the United States and race is not the same everywhere in the world. Racial categories serve particular contextual purposes depending on the society they are used in, but generally follow the base logic of the supremacy of one type of human body over all others (ordering these human bodies in a hierarchical fashion).\u201d Choosing which biological or nonbiological features to use when discussing race is always a social process (Omi and Winant 2014). Race concepts have no validity to them unless people continue to use them in their daily lives\u2014and, in the worst cases, to use them to justify racist behaviours and problematic ideas about racial difference or superiority\/inferiority. Recent work in anthropological genetics has revealed the similarities amongst humans on a molecular level and the relatively few differences that exist between populations (Omi and Winant 2014).<\/p>\n<p class=\"import-Normal\">The role of the biological anthropologist becomes crucial in the public sphere, because we may be able to debunk myths surrounding human variation and shed light, for the nonanthropologists around us, on how human variation is actually distributed worldwide (see Figure 14.1). Rooted in scientific observations, our work can help nonanthropologists recognize how common ideas about \u201crace\u201d often have no biological or genetic basis. Many anthropologists work hard to educate students on the history of where race concepts come from, why and how they last in public consciousness, and how we become more conscious of racial issues and the need to fight against racism in our societies. Throughout this chapter, I will highlight how humans cannot actually be divided into discrete \u201craces,\u201d because most traits vary on a continuous basis and human biology is, in fact, very <strong>homogenous<\/strong> compared to the greater genetic variation we observe in closely related species. Molecular anthropology, or anthropological genetics, continues to add new layers to our understanding of human biological variation and the evolutionary processes that gave rise to the contemporary patterns of human variation. The study of human variation has not always been unbiased, and thinkers and scientists have always worked in their particular sociohistorical context. For this reason, this chapter opens with a brief overview of race concepts throughout history, many of which relied on unethical and unscientific notions about different human groups.<\/p>\n<\/div>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: My Experiences as an Asian Academic<\/h2>\n<figure style=\"width: 578px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image10-4.jpg\" alt=\"Outdoor photo of this chapter\u2019s author.\" width=\"578\" height=\"433\" \/><figcaption class=\"wp-caption-text\">Figure 14.2: Michael B. C. Rivera in Hong Kong. Credit: Michael B. C. Rivera in Hong Kong original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) is under a <a class=\"rId26\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p>My name is Michael, and I am a biological anthropologist and archaeologist (Figure 14.2). What strikes me as most interesting to investigate is human biological variation today and the study of past human evolution. For instance, some of my research on ancient coastal populations has revealed positive effects of coastal living on dietary health and many unique adaptations in bones and teeth when living near rivers and beaches. I love talking to students and nonscientists about bioanthropologists\u2019 work, through teaching, science communication events, and writing book chapters like this one. I grew up in Hong Kong, a city in southern China. My father is from the Philippines and my mother is from Hong Kong, which makes me a mixed Filipino-Chinese academic. Growing up, I noticed that people came in all shapes, sizes, and colours. My life is very different now in that I have gained the expertise to explain those differences, and I feel a great responsibility to guide those new to anthropology toward their own understandings of diversity.<\/p>\n<p class=\"import-Normal\">Biological anthropology is not taught extensively in Hong Kong, so I moved to the United Kingdom to earn my bachelor\u2019s, master\u2019s, and doctorate degrees. It was fascinating to me that we could answer important questions about human variation and history using scientific methods. However, I did not have many minority academic role models to look up to while I was at university. My department was made up almost exclusively of white westerner faculty, and it was hard to imagine I could one day get a job at these western institutions. While pursuing my degrees, I also remember several instances of my research contributions being overlooked or dismissed. Sometimes professors and fellow students would make racist comments toward Asian scholars (including me) and other Black, Indigenous and researchers of colour, making us greatly uncomfortable in those spaces. When one of us would work up the courage to tell university leaders our experiences of being stereotyped, dismissed, or insulted, we received little support and were further excluded from research and teaching activities. This is a common experience for Black, Indigenous, and other people of colour who pursue biological anthropology, and we face the difficult choice between leaving the field or bearing with such unsafe spaces.<\/p>\n<p class=\"import-Normal\">It became important to me at that time to find other academics of colour with whom to share experiences and form community. I feel inspired by all my colleagues who advocate for greater representation and diversity in universities (whether they are minority academics or not). I admire many of my fellow researchers who are underrepresented and do a great job of representing minority groups through their cutting-edge research and quality teaching at the undergraduate and graduate levels. Although I no longer work in the West, it has remained my great hope that those in the West and the \u201cGlobal North\u201d will continue to improve university culture, and I support any efforts there to welcome all scholars.<\/p>\n<p class=\"import-Normal\">My current work is based in Hong Kong, where I am deeply dedicated to helping develop biological anthropology in East and Southeast Asia and promoting research from our home regions on the international scene. The study of anthropology really highlights how we share a common humanity and history. Being somebody who is \u201cmixed race\u201d and Asian likely played a key role in steering me toward the study of human variation. As this chapter hopefully shows, there is a lot to discuss about race and ethnicity regarding the discipline\u2019s history and current understandings of <strong>human diversity<\/strong>. Some scientific and technological advancements today are misused for reasons to do with money, politics, or the continuation of antiquated ideas. It is my belief, alongside many of my friends and fellow anthropologists, that science should be more about empathy toward all members of our species and contributing to the intellectual and technological nourishment of society. After speaking to many members of the public, as well as my own undergraduate students, I have received lovely messages from other individuals of colour expressing thanks and appreciation for my presence and understanding as a minority scientist and mentor figure. Anthropology needs much more diversity as well as to make room for those who have traveled different routes into the discipline. All paths taken into anthropology are valid and valuable. I would encourage everyone to study anthropology\u2014it is a field for understanding and celebrating the intricacies of human diversity.<\/p>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<h2 class=\"import-Normal\">The History of \"Race\" Concepts<\/h2>\n<h3 class=\"import-Normal\"><strong>\u201cRace\u201d in the Classical Era<\/strong><\/h3>\n<figure style=\"width: 435px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image11-8.png\" alt=\"Painting of four individuals with varied skin colors, head hair, facial hair, and clothing styles.\" width=\"435\" height=\"331\" \/><figcaption class=\"wp-caption-text\">Figure 14.3: (From left to right) Depicting a Berber (Libyan), a Nubian, an Asiatic (Levantine), and an Egyptian, copied from a mural on the tomb of Seti I. Credit: Egyptian races drawing by Heinrich von Minutoli (1820) of a mural by an unknown artist from the tomb of Seti I is in the public domain.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The earliest classification systems used to understand human variation are evidenced by ancient manuscripts, scrolls, and stone tablets recovered through archaeological, historical, and literary research. The Ancient Egyptians had the <em>Book of Gates<\/em>, dated to the New Kingdom between 1550 B.C.E. and 1077 B.C.E (Figure 14.3). In one part of this tome dedicated to depictions of the underworld, scribes used pictures and hieroglyphics to illustrate a division of Egyptian people into the four categories known to them at the time: the Aamu (Asiatics), the Nehesu (Nubians), the Reth (Egyptians), and the Themehu (Libyans). Though not rooted in any scientific basis like our current understandings of human variation today, the Ancient Egyptians believed that each of these groups were made of a distinctive category of people, distinguishable by their skin colour, place of origin, and even behavioural traits.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Another well-known early document is the Bible, where it is written that all humankind descends from one of three sons of Noah: Shem (the ancestor to all olive-skinned Asians), Japheth (the ancestor to pale-skinned Europeans), and Ham (the ancestor to darker-skinned Africans). Similar to the Ancient Egyptians, these distinctions were based on behavioural traits and skin colour. More recent work in historiography and linguistics suggest that the branches of \u201cHamites,\u201d \u201cJaphethites,\u201d and \u201cShemites\u201d may also relate to the formation of three independent language groups some time between 1000 and 3000 B.C.E. With the continued proliferation of Christianity, this concept of approximately three racial groupings lasted until the Middle Ages and spread as far across Eurasia as crusaders and missionaries ventured at the time.<\/p>\n<figure style=\"width: 309px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image9-9.png\" alt=\"Rows of organisms, with plants and animals at the bottom and humans, angels, and God at top.\" width=\"309\" height=\"449\" \/><figcaption class=\"wp-caption-text\">Figure 14.4: The Great Chain of Being from the Rhetorica Christiana by Fray Diego de Valades (1579). Credit: Great Chain of Being 2 by Didacus Valades (Diego Valades 1579) and photographed by Rhetorica Christiana (via Getty Research) is in the public domain.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">There is also the \u201cGreat Chain of Being,\u201d conceived by ancient Greek philosophers like Plato (427\u2012348 B.C.E.) and Aristotle (384\u2012322 B.C.E.). They played a key role in laying the foundations of empirical science, whereby observations of everything from animals to humans were noted with the aim of creating taxonomic categories. Aristotle describes the Great Chain of Being as a ladder along which all objects, plants, animals, humans, and celestial bodies can be mapped in an overall hierarchy (in the order of existential importance, with humans placed near the top, just beneath divine beings; see Figure 14.4). When he writes about humans, Aristotle expressed the belief that certain people are inherently (or genetically) more instinctive rulers, while others are more natural fits for the life of a worker or enslaved person. Based on research by biological anthropologists, we currently recognize that these early systems of classification and hierarchization are unhelpful in studying human biological variation. Both behavioural traits and physical traits are coded for by multiple genes each, and how we exhibit those traits based on our genetics can vary significantly even between individuals of the same population.<\/p>\n<h3 class=\"import-Normal\"><strong>\u201cRace\u201d during the Scientific Revolution<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The 1400s to 1600s saw the beginnings of the <strong>Scientific Revolution<\/strong> in Western societies, with thinkers like Nicholas Copernicus, Galileo Galilei, and Leonardo Da Vinci publishing some of their most important findings. While by no means the first or only scholars globally to use observation and experimentation to understand the world around them, early scientists living at the end of the medieval period in Europe increasingly employed more experimentation, quantification, and rational thought in their work. This is the main difference between the work of the ancient Egyptians, Romans, and Greeks and that of later scientists such as Isaac Newton and Carl Linnaeus in the 1600s and 1700s.<\/p>\n<figure style=\"width: 215px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image24-2.png\" alt=\"Historic painting of a man in 18th-century wig and garments.\" width=\"215\" height=\"259\" \/><figcaption class=\"wp-caption-text\">Figure 14.5: Carl Linnaeus. Credit: <a class=\"rId35\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Carl_von_Linn%C3%A9.png\">Carl von Linn\u00e9<\/a> by <a class=\"rId36\" href=\"https:\/\/en.wikipedia.org\/wiki\/en:Alexander_Roslin\">Alexander Roslin<\/a> (1718-1793) is in the <a class=\"rId37\" 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\">Linnaeus is the author of <em>Systema Naturae<\/em> (1758), in which he classified all plants and animals under the formalized naming system known as <strong>binomial nomenclature<\/strong> (Figure 14.5). This system is <strong>typological<\/strong>, in that organisms are placed into groups according to how they are similar or different to others under study. What was most anthropologically notable about Linnaeus\u2019s typological system was that he was one of the first to group humans with apes and monkeys, based on the anatomical similarities between humans and nonhuman primates. However, Linnaeus viewed the world in line with <strong>essentialism<\/strong>, a problematic concept that dictates that there are a unique set of characteristics that organisms of a specific kind <em>must<\/em> have and that would remove organisms from taxonomic categorizations if they lacked any of the required criteria.<\/p>\n<p>Linnaeus subdivided the human species into four varieties, with overtly racist categories based on skin colour and \u201cinherent\u201d behaviours. Some European scientists during this period were not aware of their own biases skewing their interpretations of biological variation, while others deliberately worked to shape public perceptions of human variation in ways that established \u201c<strong>otherness<\/strong>\u201d and enforced European domination and the subordination of non-European people. The conclusions and claims at which they arrived, consciously or subconsciously, often fit the times they were living through\u2014the so-called <strong>Age of Discovery<\/strong>, when the superiority of European cultures over others was a pervasive idea throughout people\u2019s social and political lives. Although much of Eurasia was linked by spice- and silk-trading routes, the European colonial period between the 1400s and 1700s was marked by many new and unfortunately violent encounters overseas (Figure 14.6). When Europeans arrived by ship on the shores of continents that were already inhabited, it was their first meeting with the Indigenous peoples of the Americas and Australasia, who looked, spoke, and behaved differently from peoples with whom they were familiar. Building on the idea of species and \u201csubspecies,\u201d natural historians of this time invented the term <em>race<\/em>, from the French <em>rasse<\/em> meaning \u201clocal strain.\u201d<\/p>\n<figure style=\"width: 421px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image25-4.png\" alt=\"Spanish explorers with their military and religious gear surrounding indigenous people.\" width=\"421\" height=\"273\" \/><figcaption class=\"wp-caption-text\">Figure 14.6: A painting depicting the colonization of the Mississippi River environs by Spaniard Hernando DeSoto in 1541 (painted in 1853 by William H. Powell). Credit: <a class=\"rId39\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Discovery_of_the_Mississippi.jpg\">Discovery of the Mississippi<\/a> by <a class=\"rId40\" href=\"https:\/\/en.wikipedia.org\/wiki\/en:William_Henry_Powell\">William Henry Powell<\/a> (photograph courtesy of <a class=\"rId41\" href=\"https:\/\/en.wikipedia.org\/wiki\/Architect_of_the_Capitol\">Architect of the Capitol<\/a>) is in the <a class=\"rId42\" href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Another scientist of the times, Johann Friedrich Blumenbach (1752\u20121840), classified humans into five races based on his observations of cranial form variation as well as skin colour. He thus dubbed the \u201coriginal\u201d form of the human cranium the \u201cCaucasian\u201d form, with the idea that the ideal climate conditions for early humans would have been in the Caucasus region near the Caspian Sea. The key insight Blumenbach presented was that human variation in any particular trait should be more accurately viewed as falling along a gradation (Figure 14.7). While some of his theories were correct according to what we observe today with more knowledge in genetics, they erroneously believed that human \u201csubspecies\u201d were \u201cdegenerated\u201d or \u201ctransformed\u201d varieties of an ancestral Caucasian or European race. According to them, the Caucasian cranial dimensions were the least changed over human evolutionary time, while the other skull forms represented geographic variants of this \u201coriginal.\u201d As will be discussed in greater detail later in this chapter, we have genetic and craniometric evidence for sub-Saharan Africa being the origin of the human species instead (see Chapter 13 on the fossil record that places the origins of modern <em>Homo sapiens<\/em> in north and east Africa). Based on work that shows how most biological characteristics are coded for by nonassociated genes, it is not reasonable to draw links between individuals\u2019 personalities and their skull shapes.<\/p>\n<figure style=\"width: 686px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image7-10.png\" alt=\"Historic drawing of five skulls.\" width=\"686\" height=\"259\" \/><figcaption class=\"wp-caption-text\">Figure 14.7: Five skull drawings representing specimens for Blumenbach\u2019s \u201cMongolian,\u201d \u201cAmerican,\u201d \u201cCaucasian,\u201d \u201cMalayan,\u201d and \u201cAethiopian\u201d races. Credit: <a class=\"rId44\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Blumenbach's_five_races.JPG\">Blumenbach's five races<\/a> by Johann Friedrich Blumenbach is in the <a class=\"rId45\" href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>. Original in the 1795 Treatise on \"De generis humani varietate nativa,\" unnumbered page at the end of the book titled \"Tab II\".<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>\u201cRace\u201d and the Dawn of Scientific Racism<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Between the 1800s and mid-1900s, and contrary to what you might expect, an increased use of scientific methods to justify racial schemes developed in scholarship. Differing from earlier views, which saw all humans as environmentally deviated from one \u201coriginal\u201d humankind, classification systems after 1800 became more <strong>polygenetic<\/strong> (viewing all people as having separate origins) rather than <strong>monogenetic<\/strong> (viewing all people as having a single origin). Instead of moving closer to our modern-day understandings of human variation, there was increased support for the notion that each race was created separately and with different attributes (intelligence, temperament, and appearance).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The 1800s were an important precursor to modern biological anthropology as we know it, given that this was when the scientific measurement of human physical features (anthropometry) truly became popularized. However, empirical studies in the 1800s pushed even further the idea that Europeans were culturally and biologically superior to others. While considered one of the pioneers of American \u201cphysical\u201d anthropology, Samuel George Morton (1799\u20121851) was a scholar who had a large role in perpetuating 1800s scientific racism. By measuring cranial size and shape, he calculated that \u201cCaucasians,\u201d on average, have greater cranial volumes than other groups, such as the Indigenous peoples of the Americas and peoples Morton referred to collectively as \u201cNegros.\u201d Today, we know that cranial size variation depends on such factors as Allen\u2019s and Bergmann\u2019s rules, which give a more likely explanation: in colder environments, it is advantageous for those living there to have larger and rounder heads because they conserve heat more effectively than more slender heads (Beals Et al. 1984). The leading figures in craniometry during the 1800s instead were linked heavily with powerful individuals and wealthy sociopolitical institutions and financial bodies. Theories in support of hierarchical racial schemes using \u201cscientific\u201d bases certainly helped continue the exploitative and unethical trafficking and enslavement of Africans between the 1500s and 1800s.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Morton went on to write in his publication <em>Crania Americana<\/em> (1839) a number of views that fit with a concept called <strong>biological determinism<\/strong>. The idea behind biological determinism is that an association exists between people\u2019s physical characteristics and their behaviour, intelligence, ability, values, and morals. If the idea is that some groups of people are essentially superior to others in cognitive ability and temperament, then it is easier to justify the unequal treatment of certain groups based on outward appearances. Another such problematic thinker was Paul Broca (1824\u20121880), after whom a region of the frontal lobe related to language use is named (Broca\u2019s area). Influenced by Morton, Broca likewise claimed that internal skull capacities could be linked with skin colour and cognitive ability. He went on to justify the European colonization of other global territories by purporting it was noble for a biologically more \u201ccivilized\u201d population to improve the \u201chumanity\u201d of more \u201cbarbaric\u201d populations. Today, these theories of Morton, Broca, and others like them are known to have no scientific basis. If we could speak with them today, they would likely try to emphasize that their conclusions were based on empirical evidence and not <em>a priori<\/em> reasoning. However, we now can clearly see that their reasoning was biased and affected by prevailing societal views at the time.<\/p>\n<h3 class=\"import-Normal\"><strong>\u201cRace\u201d and the Beginnings of Physical Anthropology<\/strong><\/h3>\n<p class=\"import-Normal\">In the early 20th century, we saw a number of new figures coming into the science of human variation and shifting the theoretical focus within. Most notably, these included Ale\u0161 Hrdli\u010dka and Franz Boas.<\/p>\n<figure style=\"width: 430px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image19-4.jpg\" alt=\"Historic photo of a middle-aged person in suit and bowtie.\" width=\"430\" height=\"558\" \/><figcaption class=\"wp-caption-text\">Figure 14.8: Ale\u0161 Hrdli\u010dka (1869\u00ad\u20121943), a Czech anthropologist who founded the American Journal of Physical Anthropology. Credit: <a class=\"rId47\" href=\"https:\/\/siarchives.si.edu\/collections\/siris_sic_10822?back=%2Fcollections%2Fsearch%3Fquery%3Dczech%26online%3Dtrue%26page%3D1%26perpage%3D10%26sort%3Drelevancy%26view%3Dlist#\">(Ales Hrdlicka) SIA2009-4246<\/a> (1903) by an unknown photographer <a class=\"rId48\" href=\"https:\/\/www.si.edu\/termsofuse\">is used for educational and non-commercial purposes as outlined by the Smithsonian.<\/a>. [<a class=\"rId49\" href=\"https:\/\/www.si.edu\/\">Smithsonian Institution<\/a> Archives, Record Unit 9521, Box 1, T. Dale Stewart Oral History Interview; and Record Unit 9528, Box 1, Henry B. Collins Oral History Interview.]<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Ale\u0161 Hrdli\u010dka (1869\u00ad\u20121943) was a Czech anthropologist who moved to the United States. In 1903, he established the physical anthropology section of the National Museum of Natural History (Figure 14.8). In 1918, he founded the <em>American Journal of Physical Anthropology<\/em>, which remains one of the foremost scientific journals disseminating bioanthropological research. As part of his work and the scope of the journal, he differentiated \u201c<strong>physical anthropology<\/strong>\u201d from other kinds of anthropology: he wrote that physical anthropology is \u201cthe study of racial anatomy, physiology, and pathology\u201d and \u201cthe study of man\u2019s variation\u201d (Hrdli\u010dka 1918). In some ways, although the scope and technological capabilities of biological anthropologists have changed significantly, Hrdli\u010dka established an area of inquiry that has continued and prospered for over a hundred years.<\/p>\n<p class=\"import-Normal\">Franz Boas (1858\u20121942) was a German American anthropologist who established the four-field anthropology system in the United States and founded the American Anthropological Association in 1902. He argued that the scientific method should be used in the study of human cultures and the comparative method for looking at human biology worldwide. One of Boas\u2019s significant contributions to biological anthropology was the study of skull dimensions with respect to race. After a long-term research project, he demonstrated how cranial form was highly dependent on cultural and environmental factors and that human behaviours were influenced primarily not by genes but by social learning. He wrote in one essay for the journal <em>Science<\/em>: \u201cWhile individuals differ, biological differences between races are small. There is no reason to believe that one race is by nature so much more intelligent, endowed with great willpower, or emotionally more stable than another, that the difference would materially influence its culture\u201d (Boas 1931:6). This conclusion directly contrasted with the theories of the past that relied on biological determinism. Biological anthropologists today have found evidence that corroborates Boas\u2019s explanations: societies do not exist on a hierarchy or gradation of \u201ccivilizedness\u201d but instead are shaped by the world around them, their demographic histories, and the interactions they have with other groups.<\/p>\n<figure style=\"width: 358px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image2-7.png\" alt=\"Black-and-white sketch of tree labeled \u201cEugenics.\u201d \" width=\"358\" height=\"275\" \/><figcaption class=\"wp-caption-text\">Figure 14.9: Logo of the Second International Exhibition of Eugenics, held in 1921. The text of the logo states: \"Eugenics is the self-direction of human evolution. Like a tree, eugenics draws its materials from many sources and organizes them into an harmonious entity.\" Credit: <a class=\"rId51\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Eugenics_congress_logo.png\">Eugenics congress logo<\/a> scanned from <a class=\"rId52\" href=\"https:\/\/en.wikipedia.org\/wiki\/Harry_H._Laughlin\">Harry H. Laughlin<\/a>, The Second International Exhibition of Eugenics held September 22 to October 22, 1921, is in the <a class=\"rId53\" 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 first half of the 1900s still involved some research that was essentialist and focused on proving racial determinism. Anthropologists like Francis Galton (1822\u20121911) and Earnest A. Hooton (1887\u20121954) created the field of <strong>eugenics<\/strong> as an attempt to formalize the social scientific study of \u201cfitness\u201d and \u201csuperiority\u201d among members of 19th-century Europe. As a way of \u201cdealing with\u201d criminals, diseased individuals, and \u201cuncivilized\u201d people, eugenicists recommended prohibiting parts of the population from being married or sterilizing these members of society so they could no longer procreate (Figure 14.9). They instead encouraged \u201creproduction in individual families with sound physiques, good mental endowments, and demonstrable social and economic capability\u201d (Hooton 1936). In the 1930s, Nazi Germany used this false idea of there being \u201cpure races\u201d to highly destructive effect. The need to be protected against admixture from \u201cunfit\u201d groups was their justification for their blatant racism and purging of citizens that fell under their subjective criteria.<\/p>\n<p class=\"import-Normal\">It should be noted that eugenicist ideas were popularly discussed and debated in many non-European contexts, as in the U.S., China, and South Africa, too. The Immigration Restriction Act of 1924 was passed in the United States, with the explicit aim of reducing the country\u2019s \u201cburden\u201d of people considered inferior by restricting immigration of eastern European Jews, Italians, Africans, Arabs, and Asians. In the early 1900s, Chinese scientists and politicians showed great interest in eugenic ideologies, which came to dictate decisions in law-making, family life, and the medical field. Noted American anthropologist Ruth Benedict wrote extensively on Japanese culture and society during and after World War II. Her essentialist portrayals of Japanese people were heavily cited in popular discourse at the time. In 1950s South Africa, interracial marriages and sexual relations were banned by law; antimiscegenation became one of the huge focuses of apartheid resistance activists in later years. We still see the continuation of eugenics-based logic today around the world\u2014in exclusionary immigration laws, cases of incarcerated prison inmates being forcibly sterilized, and the persistence of intelligence testing as a form of measuring people\u2019s \u201cfitness\u201d in a society.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Shortly after World War II and the Nazi Holocaust, the full extent of essentialist, eugenicist thinking became clear. Social constructions of race, and the notion that one can predict psychological or behavioural traits based on external appearance, had become unpopular both within and outside the discipline. It was up to those in the field of physical anthropology at the time to separate physical anthropology from race concepts that supported unscientific and socially damaging agendas. This does not mean that there are no physiological or behavioural differences between different members of the human species. However, going forward, a number of physical anthropologists saw human biological variation as more complicated than simple typologies could describe.<\/p>\n<h3 class=\"import-Normal\"><strong>\u201cThe New Physical Anthropology\u201d<\/strong><\/h3>\n<figure style=\"width: 219px\" class=\"wp-caption alignright\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image14-6.png\" alt=\"Black-and-white photo of a person with short hair in a white shirt and tie.\" width=\"219\" height=\"291\" \/><figcaption class=\"wp-caption-text\">Figure 14.10: Theodosius Dobzhansky, an important scientist who formulated the 20th-century \u201cmodern synthesis\u201d reconciling Charles Darwin\u2019s theory of evolution and Gregor Mendel\u2019s ideas on heredity. Credit: <a class=\"rId55\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Dobzhansky_no_Brasil_em_1943.jpg\">Dobzhansky no Brasil em 1943<\/a> by unknown photographer via <a class=\"rId56\" href=\"https:\/\/www.flickr.com\/photos\/celycarmo\/\">Flickr user Cely Carmo<\/a> is in the <a class=\"rId57\" href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">After 1950, focus steered away from the concept of \u201crace\u201d as a unit of variation and toward understanding why variation exists in <strong>population<\/strong><strong>s<\/strong> from an evolutionary perspective. This was outlined by those pioneering the \u201cnew physical anthropology,\u201d such as Sherwood Washburn, Theodosius Dobzhansky (Figure 14.10), and Julian Huxley, who borrowed this approach from contemporary population geneticists. Whether using genetic or phenotypic markers as the units of study, \u201cgroups\u201d or \u201cclusters\u201d of humans differentiated by these became defined as populations that differ in the frequency of some gene or genes. Anthropologists consider what \u201cmakes\u201d a population\u2014a group of individuals potentially capable of or actually interbreeding due to shared geographic proximity, language, ethnicity, culture, and\/or values. Put another way, a population is a local interbreeding group with reduced gene flow between themselves and other groups of humans. Members of the same population may be expected to share many genetic traits (and, as a result, many phenotypic traits that may or may not be visible outwardly).<\/p>\n<figure style=\"width: 240px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image15-6.png\" alt=\"Black-and-white photo of smiling person in a suit and tie in front of a building.\" width=\"240\" height=\"300\" \/><figcaption class=\"wp-caption-text\">Figure 14.11: Julian Huxley (1942). Credit: <a class=\"rId59\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Julian_Huxley_1-2.jpg\">Julian Huxley 1-2<\/a> by unknown photographer is in the <a class=\"rId60\" href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Thinking of humans in terms of populations was part of Julian Huxley\u2019s (1942) \u201cModern Synthesis\u201d\u2014so named because it helped to reconcile two fundamental principles about evolution that had not been made sense of together before (Figure 14.11). As discussed in Chapter 4, Gregor Mendel (1822\u20121884) was able to show that inheritance was mediated by discrete particles (or genes) and not blended in the offspring. However, it was difficult for some 19th-century scientists to accept this model of genetic inheritance at the time because much of biological variation appeared to be continuous and not particulate (take skin colour or height as examples). In the 1930s, it was demonstrated that traits could be polygenic and that multiple alleles could be responsible for any one phenotypic trait, thus producing the continuous variation in traits such as eye colour that we see today. Thus, Huxley\u2019s \u201cModern Synthesis\u201d outlines not only how human populations are capable of exchanging genes at the microevolutionary level but also how multiple alleles for one trait (polygenic exchanges) can cause gradual macroevolutionary changes.<\/p>\n<h2 class=\"import-Normal\">Human Variation in Biological Anthropology Today<\/h2>\n<h3 class=\"import-Normal\"><strong>Many Human Traits Are Nonconcordant<\/strong><\/h3>\n<p class=\"import-Normal\">In our studies of human (genetic) variation today, we understand most human traits to be nonconcordant (Figure 14.12). \u201c<strong>Nonconcordance<\/strong>\u201d is a term used to describe how biological traits vary independent of each other\u2014that is, they do not get inherited in a correlative manner with other genetically controlled traits. For example, if you knew an individual had genes that coded for tall height, you would not be able to predict if they are lighter-skinned or have red hair. This is different from earlier essentialist views of human variation: the idea that skin colour could predict one\u2019s brain function or even \u201ctemperament\u201d and tendencies toward criminal behaviour.<\/p>\n<figure style=\"width: 594px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image17-2-1.jpg\" alt=\"World map with human silhouettes scattered across the continents.\" width=\"594\" height=\"304\" \/><figcaption class=\"wp-caption-text\">Figure 14.12: Most human biological traits are non-concordant, meaning traits vary independently and each trait has its own pattern of distribution around the world. In this image, different colours and patterns represent trait varieties. For example, the colour and pattern of the head may represent hair colour (dark to light), but sharing dark hair with another person does not mean you will share other traits (e.g. ability to digest lactose or ABO blood type). Credit: Nonconcordance original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Katie Nelson is under a <a class=\"rId62\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Human Variation Is Clinal\/Continuous (Not Discrete)<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Frank B. Livingstone (1928\u20122005) wrote: \u201cThere are no races, only clines\u201d (1962: 279). A <strong>cline<\/strong> is a gradation in the frequency of an allele\/trait between populations living in different geographic regions. Human variation cannot be broken into discrete \u201craces,\u201d because most physical traits vary on a continuous or \u201cclinal\u201d basis. One obvious example of this is how human height does not only come in three values (\u201cshort,\u201d \u201cmedium,\u201d and \u201ctall\u201d) but instead varies across a spectrum of vertical heights achievable by humans all over the world. On the one hand, we can describe human height as exhibiting <strong>continuous <\/strong><strong>variation<\/strong>, forming a continuous pattern, but height does not vary according to where people live across the globe and does not exhibit a clinal pattern. On the other hand, skin colour variation between populations does show patterning that fits quite well on to how near or far they are from each other on a world map. This makes a trait like skin colour clinally distributed worldwide. When large numbers of genetic loci for large numbers of samples were sampled from human populations distributed worldwide during the 1960s and 1970s, the view that certain facets of human diversity were clinally distributed was further supported by genetic data.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">To study human traits that are clinally distributed, genetic tests must be performed to uncover the true frequencies of an allele or trait across a certain geographic space. One easily visible example of a clinal distribution seen worldwide is the patterning of human variation in skin colour. Whether in southern Asia, sub-Saharan Africa, or Australia, dark brown skin is found. Paler skin tones are found in higher-latitude populations such as those who have lived in areas like Europe, Siberia, and Alaska for millennia. Skin colour is easily observable as a phenotypic trait exhibiting continuous variation.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">A clinal distribution still derives from genetic inheritance; however, clines often correspond to some gradually changing environmental factor. Clinal patterns arise when selective pressures in one geographic area differ from those in another as well as when people procreate and pass on genes together with their most immediate neighbours. There are several mechanisms, selective and neutral, that can lead to the clinal distribution of an allele or a biological trait. <strong>Natural selection<\/strong> is the mechanism that produced a global cline of skin colour, whereby darker skin colour protects equatorial populations from high amounts of UV radiation; there is a transition of lessening pigmentation in individuals that reside further and further away from the tropics (Jablonski 2004; Jablonski and Chaplin 2000; see Figure 14.13). The ability and inability to digest lactose (milk sugar) among different world communities varies according to differential practices and histories of milk and dairy-product consumption (Gerbault Et al. 2011; Ingram Et al. 2009). Where malaria seems to be most prevalent as a disease stressor on human populations, a clinal gradient of increasing sickle cell anemia experience toward these regions has been studied extensively by genetic anthropologists (Luzzatto 2012). Sometimes, culturally defined mate selection based on some observable trait can lead to clinal variation between populations as well.<\/p>\n<figure style=\"width: 676px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image22-6.png\" alt=\"A global map shaded representing skin colors. \" width=\"676\" height=\"370\" \/><figcaption class=\"wp-caption-text\">Figure 14.13: A global map of skin colours shows that dark skin pigmentation is more common in areas that receive more UV radiation (near the equator and in high altitude areas). Light skin is more common at northern and southern latitudes. It is worth bearing in mind, though, that these do not tell the full story of how human skin pigmentation varies worldwide. Each region will contain populations that exhibit a range of skin tones. In this way, this map is not perfect as an illustration of skin-colour distribution. Credit: Mercator style projection map showing human skin colour according to Biasutti 1940.png by Dark Tichondrias at English Wikipedia, modified (cropped) by Tuvalkin, is under a CC BY-SA 4.0 License.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Two neutral microevolutionary processes that may produce a cline in a human allele or trait are <strong>gene flow<\/strong> and <strong>genetic drift <\/strong>(see Chapter 5). The ways in which neutral processes can produce clinal distributions is seen clearly when looking at clinal maps for different blood groups in the human ABO blood group system (Figure 14.14). For instance, scientists have identified an East-to-West cline in the distribution of the blood type <em>B<\/em> allele across Eurasia. The frequency of <em>B<\/em> allele carriers decreases gradually westward when we compare the blood groups of East and Southeast Asian populations with those in Europe. This shows how populations residing nearer to one another are more likely to interbreed and share genetic material (i.e., undergo gene flow). We also see 90%\u2012100% of native South American individuals, as well as between 70%\u201290% of Aboriginal Australian groups, carrying the <em>O<\/em> allele (Mourant, \u200b\u200bKope\u0107, and Domaniewska-Sobczak 1976). These high frequencies are likely due to random genetic drift and founder effects, in which population sizes were severely reduced by the earliest <em>O<\/em> allele-carrying individuals migrating into those areas. Over time, the <em>O<\/em> blood type has remained predominant.<\/p>\n<p class=\"import-Normal\"><img class=\"aligncenter\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image8.gif\" alt=\"World map showing varying frequencies of blood type A.\" width=\"516\" height=\"284\" \/><\/p>\n<p class=\"import-Normal\"><img class=\"aligncenter\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5.gif\" alt=\"World map shows the highest frequencies of blood type B in parts of Asia.\" width=\"518\" height=\"283\" \/><\/p>\n<figure style=\"width: 516px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image13.gif\" alt=\"World map shows the highest frequencies of blood type O.\" width=\"516\" height=\"284\" \/><figcaption class=\"wp-caption-text\">Figure 14.14a\u2013c: a. Global distribution of blood type A. b. Global distribution of blood type B. c. Global distribution of blood type O. <a href=\"https:\/\/docs.google.com\/document\/d\/1VUDKMBJYS_jNONjLxT04jQN0_z9Ua50BRN6auGSHUuU\/edit\" target=\"_blank\" rel=\"noopener\">A text description of this image is available<\/a>. Credit: a. <a class=\"rId69\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Map_of_blood_group_a.gif\">Map of blood group a<\/a> by <a class=\"rId70\" href=\"https:\/\/en.wikipedia.org\/wiki\/User:Muntuwandi\">Muntuwandi<\/a> at <a class=\"rId71\" href=\"https:\/\/en.wikipedia.org\/\">en.wikipedia<\/a> is under a <a class=\"rId72\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>. b. <a class=\"rId73\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Map_of_blood_group_b.gif\">Map of blood group b<\/a> by <a class=\"rId74\" href=\"https:\/\/en.wikipedia.org\/wiki\/User:Muntuwandi\">Muntuwandi<\/a> at <a class=\"rId75\" href=\"https:\/\/en.wikipedia.org\/\">en.wikipedia<\/a> is under a <a class=\"rId76\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>. c. <a class=\"rId77\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Map_of_blood_group_o.gif\">Map of blood group o<\/a> is based on diagrams from <a class=\"rId78\" href=\"https:\/\/anthro.palomar.edu\/vary\/vary_3.htm\">https:\/\/anthro.palomar.edu\/vary\/vary_3.htm<\/a>, reproduced from A. E. Mourant Et al. (1976), and is under a <a class=\"rId79\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Genetic Variation Is Greater Within Group than Between Groups<\/strong><\/h3>\n<figure style=\"width: 295px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image27-2.jpg\" alt=\"Two medium-sized circles labeled Asian and European largely overlap. A larger circle labeled African surrounds both. \" width=\"295\" height=\"274\" \/><figcaption class=\"wp-caption-text\">Figure 14.15: Circles represent human genetic variation. Most variants are shared among individuals on all continents. There are more variants in Africa, some of which are not found in Europe or Asia. Credit: Human Genetic Variation original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Katie Nelson is under a <a class=\"rId81\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p>One problem with race-based classifications is they relied on an erroneous idea that individuals with particular characteristics would share more similar genes with each other within a particular \u201crace\u201d and share less with individuals of other \u201craces\u201d possessing different traits and genetic makeups. However, since around 50 years ago, scientific studies have shown that the majority of human genetic differences worldwide exist <em>within<\/em> groups (or \u201craces\u201d) individually rather than <em>between<\/em> groups. Indeed, most genetic variation we see occurs in Africa, and many variants are shared among individuals on all continents (Figure 14.15).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In 2002, a landmark article by Noah Rosenberg and colleagues explored worldwide human genetic variation using an even-greater genetic data set. They used 377 highly variable markers in the human genome and sampled from 1,056 individuals representative of 52 populations. The markers chosen for study were not ones that code for any expressed genes. Because these regions of the human genome were made of unexpressed genes, we may understand these markers as neutrally derived (as opposed to selectively derived) because they do not code for functional advantages or disadvantages. These neutral genetic markers likely reflect an intricate combination of regional founder effects and population histories. Analyses of these neutral markers allowed scientists to identify that 93%\u201295% of global genetic differences, referred to as <strong>variance<\/strong>, can be accounted for by within-population differences, while only a small proportion of genetic variance (3%\u20125%) can be attributed to differences among major groups (Rosenberg Et al. 2002). This research supports the theory that distinct biological races do not exist, even though misguided concepts of race may still have real social and political consequences.<\/p>\n<h3 class=\"import-Normal\"><strong>Biological Data Fit Isolation-By-Distance and Out-of-Africa Models<\/strong><\/h3>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">One further note is that the world\u2019s population may be genetically divided into \u201cgroups,\u201d \u201csubsets,\u201d \u201cclumps,\u201d or \u201cclusters\u201d that reflect some degree of genetic similarity. These identifiable clusters reflect genetic or geographic distances\u2014either with gene flow facilitated by proximity between populations or impeded by obstacles like oceans or environmentally challenging habitats (Rosenberg Et al. 2005). Sometimes, inferred clusters using multiple genetic loci are interpreted by nongeneticists literally as \u201cancestral populations.\u201d However, it would be wrong to assume from these genetic results that highly differentiated and \u201cpure\u201d ancestral groups ever existed. These groupings reflect differences that have arisen over time due to clinal patterning, genetic drift, and\/or restricted or unrestricted gene flow (Weiss and Long 2009). The clusters identified by scientists are arbitrary and the parameters used to split up the global population into groups is subjective and dependent on the particular questions or distinctions being brought into focus (Relethford 2009).<\/p>\n<figure style=\"width: 341px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image6-7.png\" alt=\"Map of the continent of Africa with the lower two-thirds shaded.\" width=\"341\" height=\"341\" \/><figcaption class=\"wp-caption-text\">Figure 14.16: Sub-Saharan Africa (shaded dark\/green). Credit: <a class=\"rId83\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Sub-Saharan-Africa.png\">Sub-Saharan Africa<\/a> by <a class=\"rId84\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Ezeu\">Ezeu<\/a> has been designated to the <a class=\"rId85\" 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\">Additionally, research on worldwide genetic variation has shown that human variation decreases with increasing distance from sub-Saharan Africa, where there is evidence for this vast region being the geographical origin of anatomically modern humans (Liu Et al. 2006; Prugnolle, Manica, and Balloux 2005; see Figures 14.16 and 14.17). Genetic differentiation decreases in human groups the further you sample data from relative to sub-Saharan Africa because of serial founder effects (Relethford 2004). Over the course of human colonization of the rest of the world outside Africa, populations broke away in expanding waves across continents into western Asia, then Europe and eastern Asia, followed by Oceania and the Americas. As a result, founder events occurred whereby genetic variation was lost, as the colonization of each new geographical region involved a smaller number of individuals moving from the original larger population to establish a new one (Relethford 2004). The most genetic variation is found across populations residing in different parts of sub-Saharan Africa, while other current populations in places like northern Europe and the southern tip of South America exhibit some of the least genetic differentiation relative to all global populations (Campbell and Tishkoff 2008).<\/p>\n<figure style=\"width: 469px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image3-6.png\" alt=\"Two scatterplots.\" width=\"469\" height=\"516\" \/><figcaption class=\"wp-caption-text\">Figure 14.17: Comparison of the genetic distance and geographical distance between populations. In the top graph, the pattern reveals that genetic variation conforms to an Out-of-Africa model, as those populations further away from Addis Ababa in Ethiopia share a smaller number of alleles; in the bottom graph, we see the populations follow an isolation-by-distance model, as pairs of populations further apart geographically seem to have greater genetic distance (Kanitz Et al. 2018). Credit: <a class=\"rId87\" href=\"https:\/\/journals.plos.org\/plosone\/article?id=10.1371\/journal.pone.0192460\">Complex genetic patterns (figure 1)<\/a> by Kanitz Et al. (2018) is under a <a class=\"rId88\" href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/\">CC BY 4.0 License<\/a>.<\/figcaption><\/figure>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<figure style=\"width: 377px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image1-9.png\" alt=\"Two large circles connected by a small area.\" width=\"377\" height=\"309\" \/><figcaption class=\"wp-caption-text\">Figure 14.18: The founder effect is a change in a small population\u2019s gene pool due to a limited number of individuals breaking away from a parent population. Credit: <a class=\"rId90\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Bottleneck_effect.jpg\">Bottleneck effect<\/a> by <a class=\"rId91\" href=\"https:\/\/wikieducator.org\/User:Tsaneda\">Tsaneda<\/a> is under a <a class=\"rId92\" href=\"https:\/\/creativecommons.org\/licenses\/by\/3.0\/legalcode\">CC BY 3.0 License<\/a>.<\/figcaption><\/figure>\n<p>Besides fitting nicely into the <strong>Out-of-Africa model<\/strong>, worldwide human genetic variation conforms to an <strong>isolation-by-distance model<\/strong>, which predicts that genetic similarity between groups will decrease exponentially as the geographic distance between them increases (Kanitz Et al. 2018). This is because of the greater and greater restrictions to gene flow presented by geographic distance, as well as cultural and linguistic differences that occur as a result of certain degrees of isolation. Since genetic data conform to isolation-by-distance and Out-of-Africa models, these findings support the abolishment of \u201crace\u201d groupings. This research demonstrates that human variation is continuous and cannot be differentiated into geographically discrete categories. There are no \u201cinherent\u201d or \u201cinnate\u201d differences between human groups; instead, variation derives from some degree of natural selection, as well as neutral processes like <strong>population bottle-necking<\/strong> (Figure 14.18), random <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1474\"><strong>mutations<\/strong><\/a> in the DNA, genetic drift, and gene flow through between-mate interbreeding.<\/p>\n<h3 class=\"import-Normal\"><strong>Humans Have Higher Homogeneity Compared to Many Other Species<\/strong><\/h3>\n<p class=\"import-Normal\">An important fact to bear in mind is that humans are 99.9% identical to one another. This means that the apportionments of human variation discussed above only concern that tiny 0.1% of difference that exists between all humans globally. Compared to other mammalian species, including the other great apes, human variation is remarkably lower. This may be surprising given that the worldwide human population has already exceeded seven billion, and, at least on the surface level, we appear to be quite phenotypically diverse. Molecular approaches to human and primate genetics tells us that external differences are merely superficial. For a proper appreciation of human variation, we have to look at our closest relatives in the primate order and mammalian class. Compared to chimpanzees, bonobos, gorillas and other primates, humans have remarkably low average genome-wide <strong>heterogeneity<\/strong> (Osada 2005).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">When we look at chimpanzee genetic variation, it is fascinating that western, central, eastern, and Cameroonian chimpanzee groups have substantially more genetic variation between them than large global samples of human DNA (Bowden Et al. 2012; Figure 14.19). This is surprising given that all of these chimpanzee groups live relatively near one another in Africa, while measurements of human genetic variation have been conducted using samples from entirely different continents. First, geneticists suppose that this could reflect differential experiences of the founder effect between humans and chimpanzees. As it has been argued that all non-African human populations descended from a small number of anatomically modern humans who left Africa, it would be expected that all groups descended from that smaller ancestral group would be similar genetically. Second, our species is really young, given that we have only existed on the planet for around 150,000 to 300,000 years. This gave humans little time for random genetic mutations to occur as genes get passed down through genetic interbreeding and meiosis. Chimpanzees, however, have inhabited different <strong>ecological niches<\/strong>, and less interbreeding has occurred between the four chimpanzee groups over the past six to eight million years compared to the amount of gene flow that occurred between worldwide human populations (Bowden Et al. 2012).<\/p>\n<figure style=\"width: 648px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image16-7.png\" alt=\"Map of Africa showing the ranges of chimpanzees from west to east.\" width=\"648\" height=\"339\" \/><figcaption class=\"wp-caption-text\">Figure 14.19: Distribution of the genus Pan, including bonobos and the four subspecies of chimpanzee, across western and central Africa (Clee Et al. 2015). <a href=\"https:\/\/docs.google.com\/document\/d\/1VUDKMBJYS_jNONjLxT04jQN0_z9Ua50BRN6auGSHUuU\/edit\" target=\"_blank\" rel=\"noopener\">A text description of this image is available<\/a>. Credit: <a class=\"rId94\" href=\"https:\/\/bmcecolevol.biomedcentral.com\/articles\/10.1186\/s12862-014-0275-z\">Chimpanzee subspecies ranges (Figure 1)<\/a> by Clee Et al. 2015 is under a <a class=\"rId95\" 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\">Recent advances have now enabled the attainment of genetic samples from the larger family of great apes and the evaluation of genetic variation among bonobos, orangutans, and gorillas alongside that of chimpanzees and humans (Prado-Martinez Et al. 2013). Collecting such data and analyzing primate genetic variation has been important not only to elucidate how different ecological, demographic, and climatic factors have shaped our evolution but also to inform upon conservation efforts and medical research. Genes that may code for genetic susceptibilities to tropical diseases that affect multiple primates can be studied through genome-wide methods. Species differences in the genomes associated with speech, behaviour, and cognition could tell us more about how human individuals may be affected by genetically derived neurological or speech-related disorders and conditions (Prado-Martinez Et al. 2013; Staes Et al. 2017). In 2018, a great ape genomic study also reported genetic differences between chimpanzees and humans related to brain cell divisions (Kronenberg Et al. 2018). From these results, it may be inferred that cognitive or behavioural variation between humans and the great apes might relate to an increased number of cortical neurons being formed during human brain development (Kronenberg Et al. 2018). Comparative studies of human and nonhuman great ape genetic variation highlight the complex interactions of population histories, environmental changes, and natural selection between and within species. When viewed in the context of overall great ape variation, we may reconsider how variable the human species is relatively and how unjustified previous \u201crace\u201d concepts really were.<\/p>\n<h3 class=\"import-Normal\"><strong>Phenotypic Traits That Reflect Neutral Evolution<\/strong><\/h3>\n<p class=\"import-Normal\">Depending on the trait being observed, different patterns of phenotypic variation may be found within and among groups worldwide. In this subsection, some phenotypic traits that reflect the aforementioned patterns of genetic variation will be discussed.<\/p>\n<figure style=\"width: 301px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image18-4.jpg\" alt=\"Illustration of one anthropologist studying teeth, and another looking into a microscope.\" width=\"301\" height=\"301\" \/><figcaption class=\"wp-caption-text\">Figure 14.20: Contemporary anthropologists who use many types of skeletal markers have demonstrated that a majority of cranial variation occurs within populations rather than between populations and that there is a decrease in variation with distance from Africa. Credit: <a class=\"rId97\" href=\"https:\/\/img1.wsimg.com\/isteam\/ip\/0f6c1c17-41ea-4caf-b839-73c676d69f01\/DentalHeartNecklace.jpg\/:\/rs=w:1280,h:1280\">Dental Anthropologist Heart Necklace<\/a> by <a class=\"rId98\" href=\"https:\/\/anthroillustrated.com\/\">Anthro Illustrated<\/a> is under a <a class=\"rId99\" 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\">Looking beyond genetic variation briefly, recent studies have revisited biological anthropology\u2019s earlier themes of externally observable traits, such as skull shape. In the last 20 or so years, anthropologists have evaluated the level to which human cranial shape variation reflects the results from genetic markers, such as those used previously to fit against Out-of-Africa models (Relethford 2004) or those used in the apportionment of human variation between and within groups (Lewontin 1972; Rosenberg Et al. 2002). Using larger sample sizes of cranial data collected from thousands of skulls worldwide and a long list of cranial measurements, studies demonstrate a similar decrease in variation with distance from Africa and show that a majority of cranial variation occurs within populations rather than between populations (Betti Et al. 2009; Betti Et al. 2010; Manica Et al. 2007; Relethford 2001; von Cramon-Taubadel and Lycett 2008; see Figure 14.20). The greatest cranial variation is found among skulls of sub-Saharan African origin, while the least variation is found among populations inhabiting places like Tierra del Fuego at the southern tip of Argentina and Chile. While ancient and historical thinkers previously thought \u201crace\u201d categories could reasonably be determined based on skull dimensions, modern-day analyses using more informative sets of cranial traits simply show that migrations out of Africa and the relative distances between populations can explain a majority of worldwide cranial variation (Betti Et al. 2009).<\/p>\n<figure style=\"width: 250px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image21-6.png\" alt=\"Sketch of bone with bony loops at the top and coil at the bottom.\" width=\"250\" height=\"208\" \/><figcaption class=\"wp-caption-text\">Figure 14.21: Diagram of the bony labyrinth in the inner ear. Credit: <a class=\"rId101\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Bony_labyrinth.png\">Bony labyrinth<\/a> by <a class=\"rId102\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Selket\">Selket<\/a> has been designated to the <a class=\"rId103\" href=\"https:\/\/creativecommons.org\/share-your-work\/public-domain\/cc0\/\">public domain (CC0)<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">This same patterning in phenotypic variation has even been found in studies examining shape variation of the pelvis (Betti Et al. 2013; Betti Et al. 2014), the teeth (Rathmann Et al. 2017), and the human <strong>bony labyrinth<\/strong> of the ear (Ponce de Le\u00f3n Et al. 2018;Figure 14.21). The skeletal morphology of these bones still varies worldwide, but a greater proportion of that variation can still be attributed to the ways in which human populations migrated across the world and exchanged genes with those closer to them rather than those further away. Human skeletal variation in these parts of the body is continuous and nondiscrete. Given the important functions of the cranium and these other skeletal parts, we may infer that the genes that underpin their development have been relatively conserved by neutral evolutionary processes such as genetic drift and gene flow. It is also important to note that while some traits such as height, weight, cranial dimensions, and body composition are determined, in part, by genes, the underlying developmental processes behind these traits are underpinned by complex polygenic mechanisms that have led to the continuous spectrum of variation in such variables among modern-day human populations.<\/p>\n<h3 class=\"import-Normal\"><strong>Phenotypic Traits That Reflect Natural Selection<\/strong><\/h3>\n<p class=\"import-Normal\">Even though 99.9% of our DNA is the same across all humans worldwide, and many traits reflect neutral processes, there are parts of that remaining 0.1% of the human genome that code for individual and regional differences. Similarly to craniometric analyses that have been conducted in recent decades, human variation in skin colour has also been reassessed using new methods and in light of greater knowledge of biological evolution.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">New technologies allow scientists to use colour photometry to sample and quantify the visible wavelength of skin colour, in a way 19th- and 20th-century readers could not. In one report, it was found that 87.9% of global skin colour variation can be attributed to genetic differences <em>between<\/em> groups, 3.2% to those among local populations within regions, and 8.9% <em>within<\/em> local populations (Relethford 2002). This apportionment differs significantly and is the reverse situation found in the distribution of genetic differences we see when we examine genetic markers such as blood type\u2013related alleles. However, this pattern of human skin colour worldwide is not surprising, given that we now understand that past selection has occurred for darker skin near the equator and lighter skin at higher latitudes (Jablonski 2004; Jablonski and Chaplin 2000). While most genetic variation reflects neutral variation due to population migrations, geographic isolation, and restricted gene flow dynamics, some human genetic\/phenotypic variation is best explained as local adaptation to environmental conditions (i.e., selection). Given that skin colour variation is atypical compared to other genetic markers and biological traits, this, in fact, goes against earlier \u201crace\u201d typologies. This is because recent studies ironically show how so much of genetic variation relates to neutral processes, while skin colour does not. It follows that skin colour <em>cannot<\/em> be viewed as useful in making inferences about other human traits.<\/p>\n<figure style=\"width: 580px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image23-3.jpg\" alt=\"Person standing at podium in front of a screen with arms partly raised.\" width=\"580\" height=\"384\" \/><figcaption class=\"wp-caption-text\">Figure 14.22: Genomicists and biological anthropologists have dedicated efforts to improving quantitative methods of measuring hair and skin variation over the last twenty years. Dr. Nina Jablonski is one such biological anthropologist specializing in the evolution and variation of human skin pigmentation. Credit: <a class=\"rId105\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Nina_Jablonski_2016_The_Skin_of_Homo_sapiens_01_%28cropped%29.jpg\">Nina Jablonski 2016 The Skin of Homo sapiens 01 (cropped)<\/a> by <a class=\"rId106\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Ptolusque\">Ptolusque<\/a> is under a <a class=\"rId107\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\">CC BY-SA 4.0 license<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">It is also true that some populations have not been studied extensively in skin pigmentation genetics (e.g., African, Austronesian, Melanesian, Southeast Asian, Indigenous American, and Pacific Islander populations, according to Lasisi and Shriver 2018). Earlier dispersals of these populations, and their local genetic varition, will have contributed to worldwide genetic variation, inclusive of skin pigmentation variation. Gene loci we did not previously appreciate as being linked to pigmentation are now being recognized thanks to better tools, more diverse genetic samples, and more accessible datasets (Quillen Et al. 2018). Biological anthropologists look forward to further discoveries elucidating the different selective pressures and population dynamics that influence skin pigmentation evolution.<\/p>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Social Implications<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">To finish this chapter, we will consider the social, economic, political, and biological implications of poor understandings of race and the deliberate perpetuation of social and medical racism.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The Black Lives Matter movement (BLM) of 2013 began with the work of racial justice activists and community organizers Alicia Garza, Opal Tometi, and Patrissa Cullors. First incited by the murder of Trayvon Martin, a 17-year-old African American, and the acquittal of the man who shot him, BLM went on to protest against the deaths of numerous Black individuals, most of whom were killed by police officers (for example, Ahmaud Arbery was killed in February of 2020 by two white non-police officers). Some key characteristics of BLM from the start were its decentralized grassroots structure, the role of university students and social media in spreading awareness of the movement, and its embrace of other movements (e.g., climate justice, ending police brutality, feminist campaigns, queer activism, immigration reform, etc.). When George Floyd was murdered by a white police officer on May 25, 2020, the BLM gained new momentum, across 2,000-plus cities in the United States, and among many protesting against historic racism and police brutality in other contexts around the globe. Many in the biological anthropology community have responded to these events with a great dedication to working against systemic racism in society and institutions (American Association of Biological Anthropologists 2020).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">BLM continues to be an important movement, as is evidenced in the degree of community organizing, mutual aid efforts, calls for political reform, progress toward curriculum reform and equality, inclusion and diversity (EDI) work in businesses and universities, the removal of monuments honoring historical figures associated with slavery and racism, and many other important actions. Garza (2016) writes: \u201cThe reality is that race in the United States operates on a spectrum from black to white \u2026 the closer you are to white on that spectrum, the better off you are.\u201d Tometi (2016) has stated: \u201cWe need [a human rights movement that challenges systemic racism] because the global reality is that Black people are subject to all sorts of disparities in most of our challenging issues of our day. I think about climate change, and how six of the ten worst impacted nations by climate change are actually on the continent of Africa.\u201d In the words of Cullors (2016), \u201cBlack Lives Matter is our call to action. It is a tool to reimagine a world where Black people are free to exist, free to live. It is a tool for our allies to show up differently for us.\u201d We gather from their words the importance of learning from the egregious role that anthropologists have played in the past, recognizing the legacies of \u201cscientific\u201d justifications for eugenics and racism in our society today, and proactively working toward environmental and social equity.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Another major industry that engages in the quantification and interpretation of human variation is medical and clinical work (National Research Council [U.S.] Committee on Human Genome Diversity 1997). Large-scale genomic studies sampling from human populations distributed worldwide have produced detailed knowledge on variation in disease resistance or susceptibility between and within populations. Let\u2019s think about drug companies who develop medicines for Black patients particularly. The predispositions to particular diseases are higher among people of African descent than some pharmaceutical businesses have taken into account. Through targeted sampling of various world groups, clinical geneticists may also identify genetic risk factors of certain common disorders such as chronic heart disease, asthma, diabetes, autoimmune diseases, and behavioural disorders. Having an understanding of population-specific biology is crucial in the development of therapies, medicines, and vaccinations, as not all treatments may be suitable for every human, depending on their genotype. During diagnosis and treatment, it is important to have an evolutionary perspective on gene-environment relationships in patients. Typological concepts of \u201crace\u201d are not useful, given that most racial groups (whether self-identified or not) popularly recognized lack homogeneity and are, in fact, variable. <strong>Cystic fibrosis<\/strong>, for instance, occurs in all world populations but can often be underdiagnosed in populations with African ancestry because it is thought of as a \u201cwhite\u201d disease (Yudell Et al. 2016).<\/p>\n<p class=\"import-Normal\">Sociologists, law scholars, and professors of race studies have written extensively on how genetic\/technological\/medical revolutions impact people of colour. In her book, <em>Fatal Invention: How Science, Politics, and Big Business Re-create Race in the Twenty-First Century <\/em>(2013), Professor Dorothy E. Roberts writes about how technological advances have been used in resuscitating race as a biological category for dividing humans in essentialist ways (Figure 14.23). She notes how members of law enforcement have engaged in racial profiling, sometimes with the use of machine-learning and facial-recognition technologies. Ancestry-testing services also purport to tell us \u201cwhat\u201d we are and to insist that this information is \u201cwritten\u201d in our genes. Such advertising campaigns obscure the nuances of genetic variation with the primary motive of tapping into people\u2019s desire to \u201cknow themselves\u201d and driving up profits for their businesses. Commercial genetic testing reinforces the idea that genes map neatly onto race, all while generating massive stores of data in DNA databases. In Roberts\u2019s view, the myth of the biological concept of race being perpetuated in these ways undermines a just society and reproduces racial inequalities.<\/p>\n<figure style=\"width: 593px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image12-6.png\" alt=\"Book cover with two DNA strands and next to the smiling portrait of the author.\" width=\"593\" height=\"305\" \/><figcaption class=\"wp-caption-text\">Figure 14.23: Professor Dorothy E. Roberts is a sociologist, legal scholar, and expert on the relationships among technology, medicine, bioethics, policymaking, race, and racism. Credit: <a class=\"rId109\" href=\"https:\/\/kpfa.org\/episode\/talkies-august-23-2016\/\">Dorothy Roberts author of Fatal Invention<\/a> by <a class=\"rId110\" href=\"https:\/\/kpfa.org\/\">https:\/\/kpfa.org\/<\/a> is copyrighted and used with permission.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The COVID-19 pandemic has had a significant impact on the world\u2019s population, particularly people living in the economic Global South and many Black, Indigenous and communities of colour residing in the Global North. We have witnessed disproportionately high numbers of COVID-related deaths and infection cases among marginalized groups. Many immigrants and ethnic minorities in various societies have also experienced scapegoating and blame directed at them for being the source of COVID-19 spread.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">To inform us on how to interpret this current worldwide pandemic, historians and anthropologists are looking back at the lessons learned from past instances of racist medicine (discriminatory practices based on broader social discrimination) and medical racism (application of discriminatory practices justified on medical grounds). Historically, who could become doctors and medical professionals was often racialized, gendered, and class specific. This made it difficult for many to overcome prejudices against women, Black people, Indigenous individuals, or other people of colour from becoming doctors and clinical researchers in places such as South Africa and the United States. This, in turn, affects the sorts of information we know about health levels and health outcomes among these very groups. In the past decade, long-overdue attention is finally being paid to how race affects biological outcomes. For instance, researchers have focused on the negative legacies of racial discrimination and racism-induced stress on hormone (im)balances, mental health disorders, cardiovascular disease prevalence, and other health outcomes (Kuzawa and Sweet 2009; Shonkoff, Slopen, and WIlliams 2021; Williams 2018). The technology and standards of protocol in medical testing have been scrutinized (for more on how pulse oximeters were not designed with nonwhite patients in mind, for example, see Sjoding Et al. 2020). Scholars of race and medicine have also written on how illness and disease spread have often been used to perpetuate societal prejudices. This manifests as xenophobic tendencies at a societal level, such as the blaming of \u201coutgroups\u201d and increased \u201cin-group\u201d protectiveness. Overreliance on the idea that people are \u201cinherently\u201d disease carriers due to genetic or biological reasons leads to improper accounting for socioeconomic or infrastructural issues that lead to differential disease prevalence amongst minority communities. (For more on race and COVID, see Tsai 2021 as well as this textbook\u2019s Chapter 16: Contemporary Topics: Human Biology and Health.)<\/p>\n<p>Lastly, consider the changing field of forensic anthropology. In the past, forensic anthropologists ascribed <strong>ancestry<\/strong> or racial categories to sets of skeletons, reliant on the belief that different human groups will exhibit biologically \u201cdiscrete\u201d assortments so as to divide along culturally constructed categories (Sauer 1992). Now, a number of forensic anthropologists have argued that we should abandon these methods, both because it is unscientific and because it further validates and perpetuates this idea that race is biologically meaningful. As scientists, whether we affirm biological race as real has huge influence on the beliefs of members of the public, the judicolegal system, and law enforcement. Not all forensic experts agree with abandoning ancestry estimation. Some prefer to refocus on the neutral or selective <em>causes<\/em> of human biological variation, and assess how <em>probabilistic<\/em> it may be to assign bones of certain dimensions to one of several identified racial categories. These debates continue today as this textbook chapter is being written. More details on population affinity may be found in Chapter 17: Forensic Anthropology.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">It is important to remember that while it is possible to look for clues about one\u2019s ancestry or geographic origin based on skull morphology, again, the amount of distinctiveness in any given sample makes it impossible to distinguish whether a cranium belongs to one group (Relethford 2009). Individuals can vary in their skeletal dimensions by continental origin, country origin, regional origin, sex, age, environmental factors, and the time period in which they lived, making it difficult to assign individuals to particular categories in a completely meaningful way (Ousley, Jantz, and Freid 2009). When forensic reports and scientific journal articles give an estimation of ancestry, it is crucial to keep in mind that responsible assignments of ancestry will be done through robust statistical testing and stated as a probability estimate. Today, we also live in a more globalized world where a skeletal individual may have been born originally to parents of two separate traditional racial categories. In contexts of great heterogeneity within populations, this definitely adds difficulty to the work of forensic scientists and anthropologists preparing results for the courtroom (genetic testing may be comparatively more helpful in such situations).<\/p>\n<\/div>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Dig Deeper: Measuring F<sub>ST<\/sub><\/h2>\n<p class=\"import-Normal\">Richard Lewontin (1929\u2012) is a biologist and evolutionary geneticist who authored an article evaluating where the total genetic variation in humans lies. Titled \u201cThe Apportionment of Human Diversity\u201d (Lewontin 1972), the article addressed the following question: On average, how genetically similar are two randomly chosen people from the same group when compared to two randomly chosen people from different groups?<\/p>\n<p class=\"import-Normal\">Lewontin studied this problem by using genetic data. He obtained data for a large number of different human populations worldwide using 17 genetic markers (including alleles that code for various important enzymes and proteins, such as blood-group proteins). The statistical analysis he ran used a measure of human genetic differences in and among populations known as the fixation index (F<sub>ST<\/sub>).<\/p>\n<p class=\"import-Normal\">Technically, F<sub>ST<\/sub> can be defined as the proportion of total genetic variance within a <em>subpopulation<\/em> relative to the total genetic variance from an <em>entire population<\/em>. Therefore, F<sub>ST<\/sub> values range from 0 to 1 (or, sometimes you will see this stated as a percentage between 0% and 100%). The closer the F<sub>ST<\/sub> value of a population (e.g., the world\u2019s population) approaches 1, the higher the degree of genetic differentiation among subpopulations relative to the overall population (see Figure 14.24 for a detailed illustration).<\/p>\n<figure style=\"width: 561px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-5.jpg\" alt=\"Three cases each illustrate two populations with a mix of two types of alleles.\" width=\"561\" height=\"473\" \/><figcaption class=\"wp-caption-text\">Figure 14.24: This diagram shows a range of different case studies with which we may understand how FST is calculated in different populations. In Case 1, the gene pools of Populations 1 and 2 are 100% different from each other but possess 0% variation within themselves, so FST has a value of 1. When there is no genetic variation at all between two populations and 100% variation within them, as in Case 2, we see that FST is calculated as 0. When we look at Case 3, where variation between and within are some values between 0% and 100%, we will get a decimal figure for FST dependent upon how much variation there is between and within populations. It is through such comparisons of population genetic data that we may quantify the relative similarities or differences between and within populations, and we may thus speak to the nonexistence of \u201cracial groups\u201d that divide up our species into broad continental or racial categories. Credit: F<sub>ST<\/sub> original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Katie Nelson is under a <a class=\"rId112\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p>&nbsp;<\/p>\n<p>In his article, Lewontin (1972) identified that most of human genetic differences (85.4%) were found within local subpopulations (e.g., the Germans or Easter Islanders), whereas 8.3% were found between populations within continental human groups, and 6.3% were attributable to traditional \u201crace\u201d groups (e.g., \u201cCaucasian\u201d or \u201cAmerind\u201d). These findings have been important for scientifically rejecting the existence of biological races (Long and Kittles 2003).<\/p>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<h2 class=\"import-Normal\">Talking About Human Biological Variation Going Forward<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">To conclude, utilizing the term <em>races<\/em> to describe human biological variation is not accurate or productive. Using a select few hundred genetic loci, or perhaps a number of phenotypic traits, it may be possible to assign individuals to a geographic ancestry, but what constitutes a bounded genetic or geographical grouping is both arbitrary and potentially harmful owing to ethical and historical reasons. The discipline of biological anthropology has moved past typological frameworks that shoehorn continuously variable human populations into discrete and socially constructed subsets. Improvements in the number of markers, the genetic technologies used to study variation, and the number of worldwide populations sampled have led to more nuanced understandings of human variation. It is of utmost importance that scientists make the following points clear to the public:<\/p>\n<ul>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Today, we refer to different local human groups as \u201cpopulations.\u201d What constitutes a population should be carefully defined in scientific reports based on some geographical, linguistic, or cultural criteria and some degree of relativity to other closely or distantly related human groups.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Humans have significantly less genetic variation than other primates and mammals, and all human beings on Earth share 99.9% of their overall DNA. Some of the remaining 0.1% of human variation varies on a clinal or continuous basis, such as can be seen when looking at ABO blood-type <strong>polymorphisms<\/strong> worldwide.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Many biological characteristics in humans are actually determined nonconcordantly and\/or polygenically. Therefore, superiority or inferiority in human behaviour or body form cannot justifiably be linked to fixed and innate differences between groups.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">Genetic distances are correlated with geographic distances among the global human population. This is especially apparent when we consider that genetic variation is highest in sub-Saharan Africa, and average genetic heterogeneity decreases in populations further away from the African continent in accordance with the migratory history of anatomically modern <em>Homo sapiens<\/em>.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">The effects of gene flow, genetic drift, and population bottlenecking are reflected in some phenotypic traits, such as cranial shape.<\/li>\n<li class=\"import-Normal\" style=\"text-indent: 0pt\">We recognize other traits, like skin colour and lactase persistence, to be the products of many millennia of natural selective pressures influencing human biology from the external environment.<\/li>\n<\/ul>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Taken together, genetic analyses of human variation do not support 20th-century (or even-earlier) concepts of race. In discussions about human variation, these genomic results help clarify how biological variation is distributed across the human population today. Taking care to think about and debate the nature of human variation is important, because although the effects and events that produced genetic differences among groups occurred in the ancient past, sociocultural concepts about race and ethnicity continue to have real social, economic, and political consequences.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Beyond talking about variation in the university setting, it is important that teachers, researchers, and students of anthropology recognize and assume the responsibility of influencing public perspectives of human variation. Race-based classification systems were developed during the colonial era, the transatlantic trafficking of kidnapped Africans and the so-called \u201cScientific Revolution\u201d by the first \u201canthropologists\u201d and scholars of humankind\u2019s variation. Unfortunately, some of their early ideas have persisted and evolved into present-day lived realities. Some of today\u2019s politicians and socioeconomic bodies have racially charged agendas that promote racism or certain kinds of economic or racial inequalities. As anthropologists, we must acknowledge that while human \u201craces\u201d are not a biological reality, their status as a (misguided) social construction does have real consequences for many people (Antrosio 2011).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In other words, while \u201crace\u201d is a sociocultural invention, the treatment different individuals receive due to their perceived \u201crace\u201d can have significant financial, emotional, sociopolitical, and physiological costs. However\u2014and importantly assuming a \u201ccolour-blind\u201d position when it comes to the topics of \u201crace\u201d and ethnicity (especially in political discussions) is actually counterproductive, because the negative social consequences of modern \u201crace\u201d ideas could be ignored, making it harder to examine and address instances of discrimination properly (Wise 2010). Rather than shy away from these topics, we can use our scientific findings to establish socially relevant and biologically accurate ideas concerning human diversity. Today, research into genetic and phenotypic differentiation among and within various human populations continues to expand in its scope, its technological capabilities, its sample sizes, and its ethical concerns. It is thanks to such scientific work done in the past few decades that we now have a deeper understanding not only of how humans vary but also of how we are biologically a rather homogenous, intermixing world population.<\/p>\n<p>&nbsp;<\/p>\n<div class=\"textbox shaded\">\n<h2>Summary<\/h2>\n<p>Historically, most concepts of race were shaped by religious and early \u2018scientific\u2019 attempts to classify people, many of which justified inequality and racism. Today, biological anthropology emphasizes that human variation is continuous (clinal), polygenic, and shaped by both evolutionary pressures and neutral processes. Misunderstandings of race, however, continue to have serious social and medical consequences; evident in systemic racism and inequities in health care.<\/p>\n<p>Biological anthropologists now play a critical role in discrediting myths about race. By studying\u00a0 human variation, we can begin to understand evolutionary processes, adaptation, and the social implications of differences among human populations. Modern research shows that that humans are far more genetically homogenous than many other species, reinforcing a conclusion that \u2018race\u2019 is not a biological reality but a powerful social construct with real effects. Through evolutionary history and genetics, human diversity can be researched and understood, rather than than through racial categories.<\/p>\n<h2 class=\"import-Normal\">Review Questions<\/h2>\n<ul>\n<li class=\"import-Normal\">How is the genetic variation of the human species distributed worldwide?<\/li>\n<li class=\"import-Normal\">What evolutionary processes are responsible for producing genotypic\/phenotypic variation within and between human populations?<\/li>\n<li class=\"import-Normal\">Should we continue to attribute any value to \u201crace\u201d concepts older than 1950, based on our current understandings of human biological variation?<\/li>\n<li class=\"import-Normal\">How should we communicate scientific findings about human biological variation more accurately and responsibly to those outside the anthropological discipline?<\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\">Key Terms<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\"><strong>Age of Discovery<\/strong>: A period between the late 1400s and late 1700s when European explorers and ships sailed extensively across the globe in pursuit of new trading routes and territorial conquest.<\/p>\n<p class=\"import-Normal\"><strong>Ancestry<\/strong>: Biogeographical information about an individual, traced either through the study of an individual\u2019s genome, skeletal characteristics, or some other form of forensic\/archaeological evidence. Anthropologists carry out probabilistic estimates of ancestry. They attribute sets of human remains to distinctive \u201cancestral\u201d groups using careful statistical testing and should report ancestry estimations with statistical probability values.<\/p>\n<p class=\"import-Normal\"><strong>Binomial nomenclature<\/strong>: A system of naming living things developed by Linnaeus in the 1700s. It employs a scientific name made up of two italicized Latin or Greek words, with the first word capitalized and representative of an organism\u2019s genus and the second word indicating an organism\u2019s species (e.g., <em>Homo sapiens<\/em>, <em>Australopithecus afarensis<\/em>, <em>Pongo tapanuliensis<\/em>, etc.).<\/p>\n<p class=\"import-Normal\"><strong>Biological anthropology<\/strong>: A branch of study under anthropology (the study of humankind) that focuses on when and where humans and our human ancestors first originated, how we have evolved and adapted globally over time, and the reasons why we see biological variation among humans worldwide today.<\/p>\n<p class=\"import-Normal\"><strong>Biological determinism<\/strong>: The erroneous concept that an individual\u2019s behavioural characteristics are innate and determined by genes, brain size, or other physiological attributes\u2014and, notably, without the influence of social learning or the environment around the individual during development.<\/p>\n<p class=\"import-Normal\"><strong>Bony labyrinth<\/strong>: A system of interconnected canals within the auditory (ear- or hearing-related) apparatus, located in the inner ear and responsible for balance and the reception of sound waves.<\/p>\n<p class=\"import-Normal\"><strong>Cline<\/strong>: A gradient of physiological or morphological change in a single character or allele frequency among a group of species across environmental or geographical lines (e.g., skin colour varies clinally, as, over many generations, human groups living nearer the equator have adapted to have more skin pigmentation).<\/p>\n<p class=\"import-Normal\"><strong>Continuous variation<\/strong>: This term refers to variation that exists between individuals and cannot be measured using distinct categories. Instead, differences between individuals within a population in relation to one particular trait are measurable along a smooth, continuous gradient.<\/p>\n<p class=\"import-Normal\"><strong>Cystic fibrosis<\/strong>: A genetic disorder in which one defective gene causes overproduction and buildup of mucus in the lungs and other bodily organs. It is most common in northern Europeans (but also occurs in other world populations).<\/p>\n<p class=\"import-Normal\"><strong>Ecological niche<\/strong>: The position or status of an organism within its community and\/or ecosystem, resulting from the organism\u2019s structural and functional adaptations (e.g., bipedalism, omnivorous diets, lactose digestion, etc.).<\/p>\n<p class=\"import-Normal\"><strong>Essentialism<\/strong>: A belief or view that an entity, organism, or human grouping has a specific set of characteristics that are fundamentally necessary to its being and classification into definitive categories.<\/p>\n<p class=\"import-Normal\"><strong>Ethnicity<\/strong>: A term used commonly in an interchangeable way with the term <em>race<\/em>, complicated because how different people define this term depends on the qualities and characteristics they use to assign a label or identity to themselves and\/or others (which may include aspects of family background, skin colour, language(s) spoken, religion, physical proportions, behaviour and temperament, etc.).<\/p>\n<p class=\"import-Normal\"><strong>Eugenics<\/strong>: A set of beliefs and practices that involves the controlled selective breeding of human populations with the hope of improving their heritable qualities, especially through surgical procedures like sterilization and legal rulings that affect marriage rights for interracial couples.<\/p>\n<p class=\"import-Normal\"><strong>Gene flow<\/strong>: A neutral (or nonselective) evolutionary process that occurs when genes get shared between populations.<\/p>\n<p class=\"import-Normal\"><strong>Genetic drift<\/strong>: A neutral evolutionary process in which allele frequencies change from generation to generation due to random chance.<\/p>\n<p class=\"import-Normal\"><strong>Heterogeneity<\/strong>: The quality of being diverse genetically.<\/p>\n<p class=\"import-Normal\"><strong>Homog<\/strong><strong>enous<\/strong>: The quality of being uniform genetically.<\/p>\n<p class=\"import-Normal\"><strong>Human diversity<\/strong>: Human diversity is a measure of variation that may describe how many different forms of human there are, separated or clustered into groups according to some genetic, phenotypic, or cultural trait(s). The term can be applied to culture (in which case humans can be described as significantly diverse) or genetics (in which case humans are not diverse because all humans on Earth share a majority of their genes).<\/p>\n<p class=\"import-Normal\"><strong>Human variation<\/strong>: Differences in biology, physiology, body chemistry, behaviour, and culture. By measuring these differences, we understand the degrees of variation between individuals, groups, populations, or species.<\/p>\n<p class=\"import-Normal\"><strong>Isolation-by-distance model<\/strong>: A model that predicts a positive relationship between genetic distances and geographical distances between pairs of populations.<\/p>\n<p class=\"import-Normal\"><strong>Monogenetic<\/strong>: Pertaining to the idea that the origin of a species is situated in one geographic region or time (as opposed to <em>polygenetic<\/em>).<\/p>\n<p class=\"import-Normal\"><strong>Mutation<\/strong>: A gene alteration in the DNA sequence of an organism. As a random, neutral evolutionary process that occurs over the course of meiosis and early cell development, gene mutations are possible sources of variation in any given human gene pool. Genetic mutations that occur in more than 1% of a population are termed <em>polymorphisms<\/em>.<\/p>\n<p class=\"import-Normal\"><strong>Natural selection<\/strong>: An evolutionary process whereby certain traits are perpetuated through successive generations, likely owing to the advantages they give organisms in terms of chances of survival and\/or reproduction.<\/p>\n<p class=\"import-Normal\"><strong>Nonconcordance<\/strong>: The fact of genes or traits not varying with one another and instead being inherited independently.<\/p>\n<p class=\"import-Normal\"><strong>Otherness<\/strong>: In postcolonial anthropology, we now understand \u201cothering\u201d to mean any action by someone or some group that establishes a division between \u201cus\u201d and \u201cthem\u201d in relation to other individuals or populations. This could be based on linguistic or cultural differences, and it has largely been based on external characteristics throughout history.<\/p>\n<p class=\"import-Normal\"><strong>Out-of-Africa model<\/strong>: A model that suggests that all humans originate from one single group of <em>Homo sapiens<\/em> in (sub-Saharan) Africa who lived between 100,000 and 315,000 years ago and who subsequently diverged and migrated to other regions across the globe.<\/p>\n<p class=\"import-Normal\"><strong>Physical anthropology<\/strong>: This used to be the more common name given to the subdiscipline of anthropology centred upon the study of human origins, evolution and variation (also see <em>biological anthropology<\/em> above). This name for the field has gradually become less popular due to two reasons: first, it may not reflect our interests in other aspects of humankind that are not physical (such as those behavioural, cultural and spiritual), and second, using this term popular in the early decades of our field may be viewed by some as harkening back to a time when biological anthropologists conducted their work in unethical ways.<\/p>\n<p class=\"import-Normal\"><strong>Polygenetic<\/strong>: Having many different ancestries, as in older theories about human origins that involved multiple traditional groupings of humans evolving concurrently in different parts of the world before they merged into one species through interbreeding and\/or intergroup warfare. These earlier suggestions have now been overwhelmed by insurmountable evidence for a single origin of the human species in Africa (see the \u201cOut-of-Africa model\u201d).<\/p>\n<p class=\"import-Normal\"><strong>Polymorphism<\/strong>: A genetic variant within a population (caused either by a single gene or multiple genes) that occurs at a rate of over 1% among the population. Polymorphisms are responsible for variation in phenotypic traits such as blood type and skin colour.<\/p>\n<p class=\"import-Normal\"><strong>Population<\/strong>: A group of humans living in a particular geographical area, with more local interbreeding within-group than interbreeding with other groups. A limited or restricted amount of gene flow between populations can occur due to geographical, cultural, linguistic, or environmental factors.<\/p>\n<p class=\"import-Normal\"><strong>Population bottlenecking<\/strong>: An event in which genetic variation is significantly reduced owing to a sharp reduction in population size. This can occur when environmental disaster strikes or as a result of human activities (e.g., genocides or group migrations). An important example of this loss in genetic variation occurred over the first human migrations out of Africa and into other continental regions.<\/p>\n<p class=\"import-Normal\"><strong>Prejudice<\/strong>: An unjustified attitude toward an individual or group that is not based on reason, whether positive (and showing preference for one group of people over another) or negative (and resulting in harm or injury to others).<\/p>\n<p class=\"import-Normal\"><strong>Race<\/strong>: The identification of a group based on a perceived distinctiveness that makes that group more similar to each other than they are to others outside the group. This may be based on cultural differences, genetic parentage, physical characteristics, behavioural attributes, or something arbitrarily and socially constructed. As a social or demographic category, perceptions of \u201crace\u201d can have real and serious consequences for different groups of people. This is despite the fact that biological anthropologists and geneticists have demonstrated that all humans are genetically homogenous and that more differences can be found within populations than between them in the overall apportionment of human biological variation. This term is sometimes used interchangeably with <em>ethnicity<\/em>.<\/p>\n<p class=\"import-Normal\"><strong>Racism<\/strong>: Any action or belief that discriminates against someone based on perceived differences in race or ethnicity.<\/p>\n<p class=\"import-Normal\"><strong>Scientific Revolution<\/strong>: A period between the 1400s and 1600s when substantial shifts occurred in the social, technological, and philosophical sense, when a scientific method based on the collection of empirical evidence through experimentation was emphasized and inductive reasoning was used to test hypotheses and interpret their results.<\/p>\n<p class=\"import-Normal\"><strong>Typolog<\/strong><strong>ical<\/strong>: Of or describing an assortment system that relies on the interpretation of qualitative similarities or differences in the study of variation among objects or people. The categorization of cultures or human groups according to \u201crace\u201d was performed with a typological approach in the earliest practice of anthropology, but this practice has since been discredited and abandoned.<\/p>\n<p class=\"import-Normal\"><strong>Variance<\/strong>: In statistics, variance measures the dispersal of a set of data around the mean or average value.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration<strong><br \/>\n<\/strong><\/h2>\n<h3 class=\"import-Normal\"><strong>Videos<\/strong><\/h3>\n<p>American Medical Association (AMA). 2020. \u201c<a href=\"https:\/\/www.youtube.com\/watch?v=tqA3KvvscYc\" target=\"_blank\" rel=\"noopener\">Examining Race-Based Medicine<\/a>.\u201d YouTube, October 29. Accessed June 4, 2023.<\/p>\n<p>Crenshaw, Kimberl\u00e9. 2016. \u201c<a href=\"https:\/\/www.youtube.com\/watch?v=akOe5-UsQ2o\" target=\"_blank\" rel=\"noopener\">The Urgency of Intersectionality<\/a>.\u201d YouTube, December 7. Accessed June 4, 2023.<\/p>\n<p>Golash-Boza, Tanya. 2018. \u201c<a href=\"https:\/\/www.youtube.com\/watch?v=NQOimokvJXo\" target=\"_blank\" rel=\"noopener\">What Is Race? What Is Ethnicity? Is There a Difference?<\/a>.\u201d YouTube, October 28. Accessed June 4, 2023.<\/p>\n<p>Lasisi, Tina. 2020. \u201c<a href=\"https:\/\/naturalhistory.si.edu\/education\/teaching-resources\/social-studies\/webinar-how-hair-reveals-futility-race-categories\" target=\"_blank\" rel=\"noopener\">How Hair Reveals the Futility of Race Categories<\/a>.\u201d National Museum of Natural History webinar, October 21.<\/p>\n<p>Lasisi, Tina. 2022. \u201c<a href=\"https:\/\/www.youtube.com\/watch?v=_BEJvVFxKV4\" target=\"_blank\" rel=\"noopener\">Where Does My Skin Color Come From?<\/a>.\u201d PBS Terra, August 18. Accessed June 4, 2023.<\/p>\n<p>PBS Origins. 2018. \u201c<a href=\"https:\/\/www.youtube.com\/watch?v=CVxAlmAPHec\" target=\"_blank\" rel=\"noopener\">The Origin of Race in the USA<\/a>.\u201d YouTube, April 3. Accessed June 4, 2023.<\/p>\n<p>Roberts, Dorothy. 2016. \u201c<a href=\"https:\/\/www.youtube.com\/watch?v=KxLMjn4WPBY\" target=\"_blank\" rel=\"noopener\">The Problem with Race-Based Medicine<\/a>.\u201d YouTube, March 4. Accessed June 4, 2023.<\/p>\n<p>Vox. 2015. \u201c<a href=\"https:\/\/www.youtube.com\/watch?v=VnfKgffCZ7U\" target=\"_blank\" rel=\"noopener\">The Myth of Race, Debunked in 3 Minutes<\/a>.\u201d YouTube, January 13. Accessed June 4, 2023.<\/p>\n<h3 class=\"import-Normal\"><strong>Podcast Episodes<\/strong><\/h3>\n<p>Kwong, Emily, and Rebecca Ramirez. 2021. \u201c<a href=\"https:\/\/www.npr.org\/2021\/10\/05\/1043391809\/heres-a-better-way-to-talk-about-hair\" target=\"_blank\" rel=\"noopener\">Here\u2019s a Better Way to Talk about Hair: A 16 Minute Listen with Tina, Lasisi<\/a>\u201d NPR Short Wave, October 6. Accessed June 4, 2023.<\/p>\n<p>Speaking of Race. 2020. \u201c<a href=\"https:\/\/soundcloud.com\/user-88955638\/sets\/race-and-health?fbclid=IwAR2U1jdQL3XYFS5llGvYZ6uSrvPikuakmbxUZb--8voxgAMKrLbu7Ym7LGU\" target=\"_blank\" rel=\"noopener\">Race and Health series<\/a>.\u201d Speaking of Race, April 10. Accessed June 4, 2023.<\/p>\n<h3 class=\"import-Normal\"><strong>Websites<\/strong><\/h3>\n<p>Choices Program. 2023. \u201c<a href=\"https:\/\/www.choices.edu\/teaching-news-lesson\/an-interactive-timeline-black-activism-and-the-long-fight-for-racial-justice\/\" target=\"_blank\" rel=\"noopener\">An Interactive Timeline: Black Activism and the Long Fight for Racial Justice<\/a>.\u201d <em>Choices Program, Brown University<\/em> [Interactive Timeline], Updated February, 2023.<\/p>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">American Association of Biological Anthropologists. 2020. \u201c<a href=\"https:\/\/bioanth.org\/about\/position-statements\/open-letter-our-community-response-police-brutality-against-african-americans-and-call-antiracist-action\/\" target=\"_blank\" rel=\"noopener\">An Open Letter to Our Community in Response to Police Brutality against African-Americans and a Call to Antiracist Action<\/a>\u201d. <em>American Association of Biological Anthropologists<\/em>, June 10, 2020. Accessed June 4, 2023.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Antrosio, Jason. 2011. \u201c\u2018<a href=\"https:\/\/www.livinganthropologically.com\/biological-anthropology\/race-reconciled-debunks-race\/\" target=\"_blank\" rel=\"noopener\">Race Reconciled\u2019: Race Isn\u2019t Skin Color, Biology, or Genetics<\/a>.\u201d <em>Living Anthropologically <\/em>(website), June 5, 2011; updated May 20, 2020. Accessed June 4, 2023.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Beals, Kenneth L., Courtland L. Smith, Stephen M. Dodd, J. Lawrence Angel, Este Armstrong, Bennett Blumenberg, Fakhry G. Girgis, et al. 1984. \u201cBrain Size, Cranial Morphology, Climate, and Time Machines [and Comments and Reply].\u201d <em>Current Anthropology<\/em> 25 (3): 301\u2012330.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Betti, Lia, Fran\u00e7ois Balloux, Tsunehiko Hanihara, and Andrea Manica. 2010. \u201cThe Relative Role of Drift and Selection in Shaping the Human Skull.\u201d <em>American Journal of Physical Anthropology<\/em> 141 (1): 76\u201282. https:\/\/doi.org\/10.1002\/ajpa.21115.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Betti, Lia, Fran\u00e7ois Balloux, William Amos, Tsunehiko Hanihara, and Andrea Manica. 2009. \u201cDistance from Africa, Not Climate, Explains Within-Population Phenotypic Diversity in Humans.\u201d <em>Proceedings: Biological Sciences<\/em> 276 (1658): 809\u2012814. https:\/\/doi.org\/10.1098\/rspb.2008.1563.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Betti, Lia, Noreen von Cramon-Taubadel, Andrea Manica, and Stephen J. Lycett. 2013. \u201cGlobal Geometric Morphometric Analyses of the Human Pelvis Reveal Substantial Neutral Population History Effects, Even across Sexes.\u201d <em>PloS ONE<\/em> 8 (2): e55909. https:\/\/doi.org\/10.1371\/journal.pone.0055909.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Betti, Lia, Noreen von Cramon-Taubadel, Andrea Manica, and Stephen J. Lycett. 2014. \u201cThe Interaction of Neutral Evolutionary Processes with Climatically Driven Adaptive Changes in the 3D Shape of the Human Os Coxae.\u201d <em>Journal of Human Evolution<\/em> 73 (August): 64\u201274. https:\/\/doi.org\/10.1016\/j.jhevol.2014.02.021.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Boas, Franz. 1931. \u201cRace and Progress.\u201d <em>Science<\/em> 74 1905): 1\u20128.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Bowden, Rory, Tammie S. MacFie, Simon Myers, Garrett Hellenthal, Eric Nerrienet, Ronald E. 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Pearson, et al. 2018. \u201cShades of Complexity: New Perspectives on the Evolution and Genetic Architecture of Human Skin.\u201d <em>American Journal of Physical Anthropology<\/em> 168 (S67): 4\u201326.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Rathmann, Hannes, Hugo Reyes-Centeno, Silvia Ghirotto, Nicole Creanza, Tsunehiko Hanihara, and Katerina Harvati. 2017. \u201cReconstructing Human Population History from Dental Phenotypes.\u201d <em>Scientific Reports<\/em> 7: 12495. https:\/\/doi.org\/10.1038\/s41598-017-12621-y.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Relethford, John H. 2001. \u201cGlobal Analysis of Regional Differences in Craniometric Diversity and Population Substructure.\u201d <em>Human Biology<\/em> 73 (5): 629\u2012636. https:\/\/doi.org\/10.1353\/hub.2001.0073.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Relethford, John H. 2002. \u201cApportionment of Global Human Genetic Diversity Based on Craniometrics and Skin Color.\u201d <em>American Journal of Physical Anthropology<\/em> 118 (4): 393\u2012398. https:\/\/doi.org\/10.1002\/ajpa.10079.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Relethford, John H. 2004. \u201cGlobal Patterns of Isolation by Distance Based on Genetic and Morphological Data.\u201d <em>Human Biology<\/em> 76 (4): 499\u2012513. https:\/\/doi.org\/10.1353\/hub.2004.0060.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Relethford, John H. 2009. \u201cRace and Global Patterns of Phenotypic Variation.\u201d <em>American Journal of Physical Anthropology<\/em> 139 (1): 16\u201222. https:\/\/doi.org\/10.1002\/ajpa.20900.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Roberts, Dorothy. 2013. <em>Fatal Invention: How Science, Politics, and Big Business Re-Create Race in the Twenty-First Century<\/em>. New York: The New Press.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Rosenberg, Noah A., Saurabh Mahajan, Sohini Ramachandran, Chengfeng Zhao, Jonathan K. Pritchard, and Marcus W. Feldman. 2005. \u201cClines, Clusters, and the Effect of Study Design on the Inference of Human Population Structure.\u201d <em>PLoS Genetics<\/em> 1 (6): e70. https:\/\/doi.org\/10.1371 \/journal.pgen.0010070.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Rosenberg, Noah A., Jonathan K. Pritchard, James L. Weber, Howard M. Cann, Kenneth K. Kidd, Lev A. Zhivotovsky, and Marcus W. Feldman. 2002. \u201cGenetic Structure of Human Populations.\u201d <em>Science<\/em> 298 (5602): 2381\u20122385.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Sauer, Norman J. 1992. \u201cForensic Anthropology and the Concept of Race: If Races Don\u2019t Exist, Why Are Forensic Anthropologists So Good at Identifying Them?\u201d <em>Social Science and Medicine<\/em> 34 (2): 107\u2012111. https:\/\/doi.org\/10.1016\/0277-9536(92)90086-6.<\/p>\n<p class=\"import-Normal\">Shonkoff, Jack P., Natalie Slopen, and David R. Williams. 2021. \u201cEarly Childhood Adversity, Toxic Stress, and the Impacts of Racism on the Foundations of Health.\u201d <em>Annual Review of Public Health<\/em> 42: 115\u2012134.<\/p>\n<p class=\"import-Normal\">Sjoding, Michael W., Robert P. Dickson, Theodore J. Iwashyna, Steven E. Gay, and Thomas S. Valley. 2020. \u201cRacial Bias in Pulse Oximetry Measurement.\u201d <em>The New England Journal of Medicine<\/em> 383: 2477-2478.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Staes, Nicky, Chet C. Sherwood, Katharine Wright, Marc de Manuel, Elaine E. Guevara, Tomas Marques-Bonet, Michael Kr\u00fctzen, et al. 2017. \u201cFOXP2 Variation in Great Ape Populations Offers Insight into the Evolution of Communication Skills.\u201d <em>Scientific Reports<\/em> 7 (1): 1\u201210. https:\/\/doi.org\/10.1038\/s41598-017-16844-x.<\/p>\n<p class=\"import-Normal\">Tomati, Opal. 2016. \u201cAn Interview with the Founders of Black Lives Matter.\u201d TED Talks 2016, October 26\u201228. Accessed June 15, 2023. https:\/\/www.ted.com\/talks\/alicia_garza_patrisse_cullors_and_opal_tometi_an_interview_with_the_founders_of_black_lives_matter\/up-next.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Tsai, Jennifer. 2021. \u201cCOVID-19 Is Not a Story of Race, but a Record of Racism\u2014Our Scholarship Should Reflect That Reality.\u201d <em>The American Journal of Bioethics<\/em> 21 (2): 43\u201247. https:\/\/doi.org\/10.1080\/15265161.2020.1861377.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">von Cramon-Taubadel, Noreen, and Stephen J. Lycett. 2008. \u201cBrief Communication: Human Cranial Variation Fits Iterative Founder Effect Model with African Origin.\u201d <em>American Journal of Physical Anthropology<\/em> 136 (1): 108\u2012113. https:\/\/doi.org\/10.1002\/ajpa.20775.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Weiss, Kenneth M., and Jeffrey C. Long. 2009. \u201cNon-Darwinian Estimation: My Ancestors, My Genes\u2019 Ancestors.\u201d <em>Genome Research<\/em> 19: 703\u2012710. https:\/\/doi.org\/10.1101\/gr.076539.108.19.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Williams, David W. 2018. \u201cStress and the Mental Health of Populations of Color: Advancing Our Understanding of Race-related Stressors.\u201d <em>Journal of Health and Social Behavior<\/em> 59 (4): 466\u2012485.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Wise, Tim. 2010. <em>Colorblind: The Rise of Post-Racial Politics and the Retreat from Racial Equity<\/em>. San Francisco: City Lights.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Yudell, Michael, Dorothy Roberts, Rob DeSalle, and Sarah Tishkoff. 2016. \u201cTaking Race out of Human Genetics.\u201d <em>Science<\/em> 351 (6273): 564\u2012565. https:\/\/doi.org\/10.1126\/science.aac4951.<\/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_1683_2568\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_2568\"><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_1683_1698\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1698\"><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<h6>Student contributors to this chapter: Amelia Roberts, Elyse Racicot, Emmanuelle Hunter<\/h6>\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 centred 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\">Learning Objectives<\/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_1683_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. The methodology and approaches outlined below are specific to the United States. 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 colour and form to burned, fragmented bone, therefore mistakable for human remains (Figure 16.1). 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 16.1: 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_1683_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_1683_1218\"><strong>spongy (trabecular) bone<\/strong><\/a>. Knowing that most bone contains both layers helps with the macroscopic identification of bone (Figures 16.2, 16.3). 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 16.2: 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 16.3: 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_1683_1220\">osteons<\/a><\/strong>, or bone cells (Figure 16.4). 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 16.5).<\/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 16.4: 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 16.5: 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 16.6).<\/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 16.6: The compact layer of this animal bone is very thick, with almost no spongy bone visible. Compare with Figure 16.2 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_1683_1222\">epiphyses<\/a><\/strong> (ends of the bone). The epiphyses of human subadult bones are not fused to the shaft (Figure 16.7). 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 16.7: 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_1683_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_1683_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 Behaviour 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 16.8: 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 16.8). 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_1683_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_1683_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 16.9).<\/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 16.9: 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_1683_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_1683_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_1683_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 16.10).<\/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 16.10: 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 16.11). 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 16.11: 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 reanalysed, 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_1683_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. While in the U.S. 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, lip 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_1683_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_1683_1238\">phenotype <\/a><\/strong> (outward appearance) was correlated with their innate intelligence and abilities (see Chapter 14 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_1683_1240\">epiphyseal union<\/a><\/strong> and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1244\">dental development.<\/a><\/strong> Epiphyseal union<strong> (<\/strong>or <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1242\">epiphyseal fusion<\/a><\/strong>) refers to the appearance and closure of the epiphyseal plates between the primary centres of growth in a bone and the subsequent centres of growth (see Figure 16.7). Prior to complete union, the cartilaginous area between the primary and secondary centres 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 16.12: 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 16.12). 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 16.13: 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_1683_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 16.13 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 16.14: 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_1683_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 16.14 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_1683_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_1683_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_1683_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 16.15 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 16.15: 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_1683_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 colour 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 16.16: 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 16.16).<\/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 16.17: 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 16.17). 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_1683_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 16.18; 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 16.18: 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 16.19). 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 16.19: 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 16.20). 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 calibre 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 16.20: 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 colour 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_1683_1260\">antemortem<\/a> <\/strong>(before death), <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_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_1683_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 8 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 16.21: 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 colouration. Colour gradients are visible in between high and lower temperatures, with lower temperatures resulting in black colouration 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 discolouration, 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 shaded\" style=\"background: var(--lightblue)\">\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 coloured 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 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 United States 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 and 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\">Becoming a Forensic Anthropologist<\/h2>\n<p class=\"import-Normal\">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\" href=\"https:\/\/www.theabfa.org\/coursework\">https:\/\/www.theabfa.org\/coursework<\/a>.<\/p>\n<div class=\"textbox shaded\">\n<h2>Summary<\/h2>\n<p data-start=\"123\" data-end=\"728\">As a subfield of biological anthropology, forensic anthropology encompasses a wide range of methods used to better understand human remains, whether from the present or the past. Through skeletal analysis, forensic anthropologists approach the study of the deceased from multiple perspectives. For instance, they may begin by identifying whether bones are human or animal, determining whether they are modern or archaeological, and assessing whether the remains were buried alone or as part of a larger assemblage. These initial steps provide a foundation for interpreting what the remains represent.<\/p>\n<p data-start=\"730\" data-end=\"1123\">Once a clearer understanding of the remains is established, forensic anthropologists can construct a biological profile of the individual. This process involves estimating biological sex, population affinity, age at death, and stature, as well as examining unique or individualizing features. Together, these elements allow anthropologists to build a more complete picture of the deceased.<\/p>\n<p data-start=\"1125\" data-end=\"1748\">Another central responsibility of forensic anthropologists is investigating how the individual died. Trauma analysis plays a key role in this process: Was the person affected by sharp force, blunt force, projectile injuries, or thermal damage? Determining the timing of injuries (whether they occurred before, at, or after death) along with analyzing what happened to the remains afterward, helps anthropologists understand both the cause and context of death. Taphonomic changes provide additional insight into the circumstances surrounding an individual\u2019s final moments.<\/p>\n<p data-start=\"1750\" data-end=\"2492\">Working with human remains requires careful consideration and profound respect for the deceased. For this reason, strict methods and ethical guidelines are integral to the profession. Proper handling techniques ensure that human remains are treated with dignity, while ethical standards guide anthropologists in their dual role within both medical and legal systems. Because their expertise can influence the interpretation and presentation of evidence in criminal investigations, forensic anthropologists must adhere to ethical principles. These standards are outlined by the Organization of Scientific Area Committees (OSAC) for Forensic Science, administered by the National Institute of Standards and Technology (NIST).<\/p>\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 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. <em>WIREs Forensic Science<\/em>, 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. <em>Healthcare (Basel, Switzerland), 9<\/em>(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. <em>Cureus<\/em>. <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_1683_1699\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1699\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<div class=\"textbox\">\n<p>Learn more about <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/about-the-authors\/\">authors and editors<\/a>,<a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/student-contributors\/\"> Hess' student contributions<\/a>, and <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/versioning-history\/\">versioning history<\/a>.<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\">Learning Objectives<\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li>Describe the major transitions in patterns of disease that have occurred throughout human evolution.<\/li>\n<li>Describe what is meant by a \u201cmismatch\u201d between our evolved biology and contemporary lifestyles and how this is reflected in modern disease patterns.<\/li>\n<li>Explain how the human stress response can positively and negatively have an impact on health.<\/li>\n<li>Explain what a \u201csyndemic\u201d is and why the COVID-19 pandemic represents one.<\/li>\n<li>Describe the ways institutionalized racism and bias in the medical field contributed to different rates of exposure, differential treatment, morbidity, and mortality from COVID-19 for different ethnic groups in the United States.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p>When was the last time you needed to do research for an upcoming paper? I bet you started by looking for information online. How did you go about your search? Which websites looked promising? Which ones did not entice you to click past the home page? Once you found one you thought might be useful, how much time did you spend searching for information? At what point did you decide to leave that site and move on? I would wager money that you never once thought your behaviour had anything to do with human evolution, but it does.<\/p>\n<p class=\"import-Normal\">Although we may not often stop to think about it, our evolutionary past is reflected in many aspects of modern life. The ways we \u201cforage\u201d for information on the internet mimics the ways we once foraged for food during our several-million-year history as hunter-gatherers (Chin Et al. 2015). Humans are visual hunters (Lieberman 2006). We practice optimal foraging strategy, meaning we make decisions based on energy return for investment (McElroy &amp; Townsend 2009). When we search for information online, we locate a \u201cpatch,\u201d in this case a website or research article, then quickly scan the contents to discern how much of it is useful to us. Like our hominin ancestors, we spend more time in \u201cpatches\u201d with abundant resources and abandon sites quickly once we have exhausted the available goods. As with internet searches, our evolutionary past is also reflected in the kinds of landscapes we find appealing, the foods that taste good to us, why we break a sweat at the gym, and why we have to go to the gym at all (Bogin 1991; Dutton 2009; Lieberman 2015). Many of the health problems facing humans in the 21st century also have their beginnings in the millions of years we roamed the earth as foragers.<\/p>\n<h2 class=\"import-Normal\">Preagricultural Humans<\/h2>\n<h3 class=\"import-Normal\"><strong>Diet<\/strong><\/h3>\n<p class=\"import-Normal\">Humans may be the species with the longest list of nutritional requirements (Bogin 1991). This is due to the fact that we evolved in environments where there was a high diversity of edible species but low densities of any given species. <em>Homo sapiens sapiens<\/em> require 45\u201350 essential nutrients for growth, maintenance, and repair of cells and tissues. These include protein, carbohydrates, fats, vitamins, minerals, and water. As a species, we are (or were) physically active with high metabolic demands. We are also <strong>omnivorous<\/strong> and evolved to choose foods that are dense in essential nutrients. One of the ways we identified high-calorie resources in our evolutionary past was through taste, and it is no accident that humans find sweet, salty, fatty foods appealing.<\/p>\n<p class=\"import-Normal\">The human predisposition toward sugar, salt, and fat is innate (Farb &amp; Armelagos 1980). Receptors for sweetness are found in every one of our mouth\u2019s 10,000 taste buds (Moss 2013). Preference for sweet makes sense in an ancestral environment where sweetness signaled high-value resources like ripe fruits. Likewise, \u201cthe long evolutionary path from sea-dwelling creatures to modern humans has given us salty body fluids, the exact salinity of which must be maintained\u201d (Farb &amp; Armelagos 1980), drawing us to salty-tasting things. Cravings for fat are also inborn, with some archaeological evidence suggesting that hominins collected animal bones for their fatty marrow, which contains two essential fatty acids necessary for brain development (Richards 2002), rather than for any meat remaining on the surface (Bogin 1991).<\/p>\n<p class=\"import-Normal\">Bioarchaeological studies indicate Paleolithic peoples ate a wider variety of foods than many people eat today (Armelagos Et al. 2005; Bogin 1991; Larsen 2014; Marciniak &amp; Perry 2017). Foragers took in more protein, less fat, much more fibre, and far less sodium than modern humans typically do (Eaton, Konner, &amp; Shostak 1988). Changes in tooth and intestinal morphology illustrate that animal products were an important part of human diets from the time of <em>Homo erectus<\/em> onward (Baltic &amp; Boskovic 2015; Richards 2002; Wrangham 2009). Once cooking became established, it opened up a wider variety of both plant and animal resources to humans. However, the protein, carbohydrates, and fats preagricultural peoples ate were much different from those we eat today. Wild game lacked the antibiotics, growth hormones, and high levels of cholesterol and saturated fat associated with industrialized meat production today (Walker Et al. 2005). Wild game was also protein dense, providing only 50% of energy as fat (Lucock Et al. 2014). The ways meat is prepared and eaten today also have implications for disease.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Meats cooked well done over high heat and\/or over an open flame, including hamburgers and barbecued meats, are highly carcinogenic due to compounds formed during the cooking process (Trafialek &amp; Kolanowski 2014). Processed meats that have been preserved by smoking, curing, salting, or by adding chemical preservatives such as sodium nitrite (e.g., ham, bacon, pastrami, salami, and beef jerky) have been linked to cancers of the colon, lung, and prostate (Abid, Cross, &amp; Sinha 2014; Figure 17.1). Nitrites\/nitrates have additionally been linked to cancers of the ovaries, stomach, esophagus, bladder, pancreas, and thyroid (Abid, Cross &amp; Sinha 2014). In addition, studies analyzing the diets of 103,000 Americans for up to 16 years indicate that those who ate grilled, broiled, or roasted meats more than 15 times per month were 17% more likely to develop high blood pressure than those who ate meat fewer than four times per month, and participants who preferred their meats well done were 15% more likely to suffer from <strong>hypertension<\/strong> (high blood pressure) than those who ate their meats rare (Liu 2018). A previous study of the same cohort indicated \u201cindependent of consumption amount, open-flame and\/or high-temperature cooking for both red meat and chicken is associated with an increased risk of type 2 diabetes among adults who consume animal flesh regularly\u201d (Liu Et al. 2018). Although meat has been argued to be crucial to cognitive and physical development among hominins (Wrangham 2009), there has been an evolutionary trade-off between the ability to preserve protein through cooking and the health risks of cooked meat and chemical preservatives.<\/p>\n<figure style=\"width: 343px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/06\/image1-6.png\" alt=\"Consecutive circles outline categories of cancer risk with images of processed meats and red meat.\" width=\"343\" height=\"424\" \/><figcaption class=\"wp-caption-text\">Figure 17.1: Positive associations have been observed between meat consumption and some types of cancer. The International Agency for Research on Cancer (2018) categorized four groupings of cancer risk. The first group is labeled \"causes cancer\", and the second group \"probably causes cancer\". Group 1 includes processed meats such as bacon, salami and hot dogs. Group 2A includes red meat such as beef, pork and lamb. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-11\/\">Carcinogenic Meats (Figure 16.1)<\/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>. [Includes <a href=\"https:\/\/pngimg.com\/download\/10217\">Hot dog PNG image<\/a> by unknown, <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0<\/a>; <a href=\"https:\/\/www.publicdomainpictures.net\/en\/view-image.php?image=109418&amp;picture=rasher-of-bacon\">Rasher of Bacon<\/a> by unknown, <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/\">public domain (CC0)<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Salami_aka.jpg\">Salami aka<\/a> by Andr\u00e9 Karwath <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Aka\">Aka<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.5\/legalcode\">CC BY-SA 2.5<\/a>; <a href=\"https:\/\/pngimg.com\/download\/2127\">Cow PNG image<\/a> by unknown, <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0<\/a>; <a href=\"https:\/\/pngimg.com\/download\/2184\">sheep PNG image<\/a> by unknown, <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0<\/a>; <a href=\"https:\/\/www.publicdomainpictures.net\/en\/view-image.php?image=55516&amp;picture=pig-on-white-background\">Pig on white background<\/a> by unknown, <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/\">public domain (CC0)<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Although carbohydrates represent half of the diet on average for both ancient foragers and modern humans, the types of carbohydrates consumed are very different. Ancient foragers ate fresh fruits, vegetables, grasses, legumes, and tubers, rather than the processed carbohydrates common in industrialized economies (Moss 2013). Their diets also lacked the refined white sugar and corn syrup found in many modern foods that contribute to the development of diabetes (Pontzer Et al. 2012).<\/p>\n<h3 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Physical Activity Patterns<\/strong><\/h3>\n<p class=\"import-Normal\">How do we know how active our ancestors were? Hominin morphology and physiology provide us with clues. Adaptations to heat discussed in Chapter 10 evolved in response to the need for physical exertion in the heat of the day in equatorial Africa. Human adaptations for preventing hyperthermia (overheating) suggest an evolutionary history of regular, strenuous physical activity. Research with modern foraging populations also offers clues to ancient activity patterns. Although subject to sampling biases and marginal environments (Marlowe 2005), modern foragers provide the only direct observations of human behaviour in the absence of agriculture (Lee 2013). From such studies, we know foragers cover greater distances in single-day foraging bouts than other living primates, and these treks require high levels of cardiovascular endurance (Raichlen &amp; Alexander 2014). Recent research with the Hadza in Tanzania indicates they walk up to 11 kilometres (6.8 miles) daily while hunting and gathering (Pontzer Et al. 2012), engaging in moderate-to-vigorous physical activity for over two hours each day\u2014meeting the U.S. government\u2019s weekly requirements for physical activity in just two days (Raichlen Et al. 2016; Figure 17.2). The fact that humans were physically active in our evolutionary past is also supported by the fact that regular physical exercise has been shown to be protective against a variety of health conditions found in modern humans, including <strong>cardiovascular disease<\/strong> (Raichlen &amp; Alexander 2014) and Alzheimer\u2019s dementia (Mandsager, Harb, &amp; Cremer 2018), even in the presence of brain pathologies indicative of cognitive decline (Buchman Et al. 2019).<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 624px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image10-7.png\" alt=\"Two hunters with dogs and bows walk in a savannah. \" width=\"624\" height=\"417\" \/><figcaption class=\"wp-caption-text\">Figure 17.2: Hadza foragers hunting on foot. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Hadazbe_returning_from_hunt.jpg\">Hadazbe returning from hunt<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/7177420@N03\">Andreas Lederer<\/a> has been modified (cropped) and is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\">CC BY 2.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Infectious Disease<\/strong><\/h3>\n<p class=\"import-Normal\">Population size and density remained low throughout the Paleolithic, by some estimates only 0.4 inhabitants per square kilometre (McClellan &amp; Dorn 2006). This limited <strong>morbidity<\/strong> and <strong>mortality <\/strong>from infectious diseases, which sometimes require large populations to sustain epidemics. Our earliest ancestors had primarily two types of infections with which to contend (Armelagos 1990). The first were organisms that adapted to our prehominin ancestors and have been problems ever since. Examples include head lice, pinworms, and yaws. A second set of diseases were <strong>zoonoses<\/strong>, diseases that originate in animals and mutate into a form infectious to humans. One example is the Human Immunodeficiency Virus (HIV) that originated in nonhuman primates and was likely passed to humans through the butchering of hunted primates for food (Sharp &amp; Hahn 2011). Zoonoses that could have infected ancient hunter-gatherers include tetanus and <strong>vector-borne diseases<\/strong> transmitted by flies, mosquitoes, fleas, midges, ticks, and the like. Many of these diseases are slow acting, chronic, or latent, meaning they can last for weeks, months, or even decades, causing low levels of sickness and allowing victims to infect others over long periods of time. Survival or cure does not result in lasting immunity, with survivors returning to the pool of potential victims. Such diseases often survive in animal reservoirs, reinfecting humans again and again (Wolfe Et al. 2012). A study of bloodsucking insects preserved in samples of amber dating from 15 to 100 million years ago indicates that they carried microorganisms that today cause diseases such as river blindness, typhus, Lyme disease, and malaria (Poinar 2018). Such diseases may have been infecting humans throughout our history and may have had significant impacts on small foraging communities because they more often infected adults, who provided the food supply (Armelagos Et al. 2005).<\/p>\n<h3 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Health<\/strong><\/h3>\n<p class=\"import-Normal\">Given their diets, levels of physical activity, and low population densities, nomadic preagricultural humans were likely in better health than many modern populations. This assertion is supported by comparative research conducted with modern foraging and industrialized populations. Measures of health taken from 20th-century foraging populations demonstrate excellent aerobic capacity, as measured by oxygen uptake during exertion, and low body-fat percentages, with <strong>triceps skinfold measurements<\/strong> half those of white Canadians and Americans. Serum cholesterol levels were also low, and markers for diabetes, hypertension, and cardiovascular disease were missing among them (Eaton, Konner, &amp; Shostak 1988; Raichlen Et al<em>. <\/em>2016).<\/p>\n<h2 class=\"import-Normal\">Health Consequences of the Transition to Agriculture and Animal Domestication<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The shift from foraging to food production occurred relatively recently in our evolutionary history (Larsen 2014), and there are indications our biology has not yet caught up (Pritchard 2010). Beginning around 12,000 BCE in several parts of the globe, humans began to move toward a diet based on domesticated plants and animals (Armelagos Et al. 2005). This involved manipulating the natural landscape to facilitate intensive food production, including the clearing of forest and construction of wells, irrigation canals, and ditches, exposing humans to water-borne illnesses and parasites and attracting mosquitos and other vectors of disease to human settlements. The heavy, repetitive physical labour of early agricultural production resulted in negative impacts on articular joints, including <strong>osteoarthritis<\/strong> (Larsen 2014). At the same time, nutritional diversity became restricted, focused on major cereal crops that continue to dominate agricultural production today, including corn, wheat, and rice (Jain 2012). This represented a major shift in diet from a wide variety of plant and animal foods to dependence on starchy carbohydrates, leading to increases in dental caries (cavities), reductions in stature and growth rates, and nutritional deficiencies (Larsen 2014). Domesticated animals added new foods to the human diet, including meat that was higher in fat and cholesterol than wild game as well as dairy products (Lucock Et al. 2014). Agriculture provided the means to produce a storable surplus for the first time in human history, leading to the beginnings of economic inequality (see Chapter 13). Social hierarchies led to the unequal distribution of resources, concentrating infectious disease among the poor and malnourished (Zuckerman Et al. 2014), a situation that continues to plague humanity today (Marmot 2005).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Sedentism<\/strong> and a rise in population density accompanied the move to agriculture, increasing the risk of infectious disease. Agriculture often provided enough calories, if not enough nutrition, to increase fertility. Although diets were worse and people unhealthier, populations continued to grow, even in the midst of high levels of child and maternal mortality and short life expectancies (Omran 2005). Hygiene became an issue as large settlements increased the problem of removing human waste and providing uncontaminated water (Armelagos Et al. 2005). Domesticated animals provided reservoirs of zoonotic pathogens, which affected farmers more than foragers, as farmers were in closer proximity to their animals on a daily basis (Marciniak &amp; Perry 2017). Many of these diseases became major killers of humankind, including influenza, tuberculosis, malaria, plague, syphilis, and smallpox, functioning as selective pressures in and of themselves (Cooling 2015). As these diseases encountered large human populations, they caused major epidemics that traveled along newly established routes for trade, warfare, and colonization.<\/p>\n<h2 class=\"import-Normal\">Epidemiological Transitions<\/h2>\n<p class=\"import-Normal\">Changes in diet and physical-activity patterns, population densities, and exposure to zoonoses associated with agriculture resulted in an epidemiological transition, a shift in the causes of morbidity (sickness) and mortality (death) among humankind (Omran 2005). The first epidemiological transition from foraging to food production resulted in increases in dental caries (see Chapter 13), nutritional deficiencies, infectious disease, and skeletal conditions like osteoarthritis, as well as decreases in growth and height (Larsen 2014). A second epidemiological transition occurred following the Industrial Revolution in Western Europe and the United States when improved standards of living, hygiene, and nutrition minimized the effects of infectious disease, after which people began to experience higher rates of <strong>noncommunicable diseases<\/strong>, such as <strong>cancer<\/strong>, heart disease, and diabetes due to the changes in lifestyle, diet, and activity levels that are the subject of this chapter (Omran 2005). With the addition of immunizations and other public health initiatives, modified forms of this transition remain ongoing in many low- and middle-income countries (Zuckerman Et al. 2014), with several now facing a <strong>\u201cdouble burden\u201d <\/strong>of disease, with poor, often rural, populations struggling with infectious diseases due to malnutrition, lack of sanitation, and access to health care, while more affluent citizens are victims of chronic illnesses. A third epidemiological transition is now underway as infectious diseases, including new, re-emergent, and multidrug-resistant diseases, have once again become major health concerns (Harper &amp; Armelagos 2010; Zuckerman Et al. 2014). These include COVID-19, Ebola, HIV\/AIDS, tuberculosis, malaria, dengue, Lyme disease, and West Nile virus\u2014all zoonoses that initially spread to humans through contact with animals.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Patterns of morbidity and mortality continue to shift across the globe. As with the first epidemiological transition resulting from the adoption of large-scale agriculture, such shifts can be the direct, if unintended, result of human interactions with the environment. For example, there has been a rise in chronic inflammatory diseases (CIDs) in developed countries (Versini Et al<em>.<\/em> 2015). This includes increased rates of allergic conditions like asthma and autoimmune diseases like rheumatoid arthritis, multiple sclerosis, Crohn\u2019s disease, and inflammatory bowel disease. This has coincided with the decrease in infectious disease associated with the second epidemiological transition, and the two are related. The \u201chygiene hypothesis\u201d postulates the rise in CIDs is a result of limited exposure to nonlethal environmental pathogens in utero and early childhood (Zuckerman &amp; Armelagos 2014). Modern human societies have become so sanitized that we are no longer exposed to microorganisms that stimulate the development of a healthy immune system (Versini Et al. 2015). \u201cIn effect, the lifestyle changes\u2014sanitary improvements, pasteurization, use of antibiotics, and improved hygiene\u2014that contributed to the second transition may have produced a substantial trade-off, with developed nations exchanging a high burden of infectious disease for a higher burden of CIDs\u201d (Zuckerman Et al. 2014).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The third epidemiological transition, the re-emergence of infectious disease, reflects the continuing relationship between humans, animals, and pathogens. Over 60% of <strong>e<\/strong><strong>merging infectious diseases (EIDs)<\/strong> since 1940 have been of zoonotic origin, with over 70% stemming from human contact with wildlife (Jones Et al. 2008), including COVID-19. The crossover of COVID to humans is believed to have involved transmission from bats to an intermediate species then to humans, with infected humans then passing it to other humans in a wet market in Wuhan, China in late 2019 (Worobey Et al. 2022). Two COVID variants, representing two distinct crossover events from animals to humans, were circulating in the market by February 2020. Similarly, the global bushmeat trade currently devastating Africa\u2019s wildlife is a continuing source of Ebola infection (Asher 2017), as well as the original source of HIV and viruses related to leukaemia and lymphoma among humans (Zuckerman Et al. 2014). New strains of avian (bird) flu, some with mortality rates as high as 60% among humans (WHO n.d.), are transmitted to humans through poultry production and contact with wild birds (Davis 2005). Lastly, the use of antibiotics in commercial meat production is directly related to the rise of drug-resistant strains of previously controlled infectious diseases. An estimated 80% of antibiotics in the U.S. are used to promote growth and prevent infection in livestock, and drug-resistant bacteria from these animals are transmitted to humans through meat consumption (Ventola 2015).<\/p>\n<figure style=\"width: 299px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image13-7.png\" alt=\"Aerial photograph of a flooded city.\" width=\"299\" height=\"399\" \/><figcaption class=\"wp-caption-text\">Figure 17.3: Flooding in Sindh, Pakistan, in 2022. Credit: <a href=\"https:\/\/www.flickr.com\/photos\/193804179@N08\/52331043544\/\">Flood in Pakistan<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/193804179@N08\/\">Ali Hyder Junejo<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\">CC BY 2.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">A fourth epidemiological transition is currently underway in which some parts of the globe are suffering from a <strong>\u201ctriple burden\u201d<\/strong> of infectious and chronic diseases combined with injuries and diseases related to intensifying globalization, urbanization, deforestation, and climate change (Karn &amp; Sharma 2021). Massive flooding in Pakistan in 2022 (Figure 17.3) will serve to illustrate the concept. Following a severe heat wave in June 2022, Pakistan experienced extremely heavy seasonal monsoon rains, in some provinces 700% above normal. Combined with water flow from melting glaciers, this caused the worst flooding in the country\u2019s history, putting one third of the nation under water (Sheerazi 2022). The heat wave, glacial melt, and extreme rainfall were all attributable to global climate change, inflicting destruction and disease on Pakistan, which produces less than 1% of total global carbon emissions (Government of Pakistan 2021).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">As a direct result of the flooding, infrastructure, including roads, homes, and bridges, was destroyed, and 1,700 people died, nearly 13,000 were injured, and over 33 million were displaced. In addition to their initial injuries and trauma, displaced people lacked food, health care, safe water, and basic sanitation, leading to starvation and exposure to infectious diseases like malaria and dengue fever, as well as skin conditions like scabies, caused by mites. Pakistan also has a poverty rate of 30\u201340%, contributing to already-high rates of HIV, tuberculosis, and hepatitis. At the same time, the leading causes of death are heart disease, cancer, lower respiratory diseases, and stroke (CDC n.d), all chronic conditions.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">These examples illustrate continuing interaction between humans, our evolved biology, and the physical and cultural environments in which we live. The remainder of this chapter will focus on selected diseases and the social, cultural, and environmental factors that contribute to their <strong>prevalence<\/strong> in modern, industrialized economies. We begin with the health condition that affects all of the others\u2014<strong>obesity<\/strong>.<\/p>\n<h2 class=\"import-Normal\">Obesity<\/h2>\n<p class=\"import-Normal\">According to the World Health Organization (2017), 1.9 billion of the world\u2019s people are overweight and 650 million of these are obese. In the United States, 70% of Americans are overweight, and 40% of these meet the criteria for obesity. For the first time in human history, most of the world's population lives in countries where overweight and obesity kill more people than hunger (Figure 17.4). Improvements in public health and food production have allowed a greater number of people to live past childhood and to have enough to eat. This does not include everyone. Many people still struggle with poverty, hunger, and disease, even in the wealthiest of nations, including the United States. On a global scale, however, many people not only have enough food to survive but also to gain weight\u2014enough extra weight to cause health problems.<\/p>\n<p><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image4-5.png\" alt=\"&quot;Bar\" \/><\/p>\n<p class=\"import-Normal\">Although studies show differences in daily energy expenditure between foraging and farming populations compared with industrialized peoples, the major contributor to obesity in Western populations is energy intake (Pontzer Et al. 2012). Many people not only eat too much but too much of the wrong things. Biological anthropologist Leslie Lieberman (2006) argues that contemporary humans continue to rely on cues from foraging strategies of our evolutionary past that are now counterproductive in the <strong>obesogenic<\/strong> environments in which we now live.<\/p>\n<p class=\"import-Normal\">Examine your own eating habits in the context of how humans once hunted and gathered. Humans once relied on visual cues to find food, often traveled long distances to obtain it, then transported it back to our home base. There they may have had to process it by hand to render it edible. Think of how much less energy it takes to find food now. If we have the financial resources, we can acquire big energy payoffs by simply sitting at home and using an app on our mobile phone to place an order for delivery. And, voila! High-calorie (if not highly nutritious) food arrives at our door. Should we venture out for food, search time is reduced by signage and advertising directing us toward high-density \u201cpatches\u201d where food is available 24 hours a day. These include vending machines, gas stations, and fast-food outlets. Travel time is minimal and little human energy is used in the process (Lieberman 2006).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Foods are also prepackaged and prepared in ways that allow us to eat large quantities quickly. Think French fries or chicken nuggets, which we can easily eat with our hands while doing other things, like driving or watching television, rendering eating mindless and allowing us to take in food faster than our <strong>endocrine systems<\/strong> can tell us we are getting full. Modern \u201cpatches\u201d offer low-fibre, calorie-dense resources, which allow us to eat larger quantities, a problem already encouraged by larger portion sizes (Lieberman 2006). Processed foods are also engineered to appeal to human preferences for sweet tastes and fatty, creamy textures (Moss 2013). Remember from earlier chapters that natural selection favoured depth perception, colour vision, grasping hands, and coordinated eye-hand movements as general primate traits. Advertising and packaging now use our colour vision against us, attracting us to products that have little nutritional value but that play to our evolutionary predisposition toward variety. Remember those 50 different nutrients we require? That variety is now presented to us in the form of 55 different flavours of Oreo cookies (Cer\u00f3n 2017), which we take out of the package and dip in milk using our hand-eye coordination and depth perception.<\/p>\n<p class=\"import-Normal\">Even if we are ostensibly eating the same things our ancestors did, these foods are nothing alike. Take potatoes, for example. One medium-sized, plain, baked potato is a healthy food, especially if we eat the skin too. It contains 110 calories, 0 grams of fat, 26 grams of carbohydrates, and 3 grams of protein, plus 30% of the U.S. Recommended Daily Allowance (RDA) of vitamin C, 10% of vitamin B6, 15% of potassium, and no sodium (<a href=\"https:\/\/potatogoodness.com\/\" target=\"_blank\" rel=\"noopener\">Potato USA<\/a>). In contrast, a medium order of McDonald\u2019s fries, which takes the potato and adds salt and fat, contains 340 calories, 16 grams of fat, 44 grams of carbohydrates, 4 grams of protein, and 230 mg of sodium (<a href=\"https:\/\/www.mcdonalds.com\" target=\"_blank\" rel=\"noopener\">McDonalds<\/a>). Potato chips take food processing to a whole new level, removing even more nutrition and adding a host of additional ingredients, including oils, preservatives, and artificial flavourings and colours (Moss 2013). Take Ruffles Loaded Bacon and Cheddar Potato Skins Potato Chips as an example (St. Pierre 2018). The number of ingredients increases from one to 11 to 35 as we move from the potato to the potato chip, moving further from nature with each step (Figure 17.5). It should be noted that the nutritional information for the potato chips is based on a serving size of 11 chips, an amount likely smaller than many people eat. Many sweet, fatty, salty foods like fries and chips are cheap, which is why many people choose to eat them (Moss 2013). The price of a medium-sized order of McDonald\u2019s fries as of this writing is US$1.79, and the potato chips are $2.98 for an 8.5-ounce bag. A single potato prewrapped for microwaving is available in many supermarkets for US$1.99 but requires access to a microwave and eating utensils, making it less convenient.<\/p>\n<\/div>\n<div style=\"text-align: left\">\n<table class=\"grid\" style=\"border-collapse: collapse;width: 100%;height: 197px\" border=\"0\">\n<caption>Figure 17.5: The potato in three modern forms. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-11\/\">The potato in three modern forms (Figure 16.4)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Joylin Namie and 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>. [Includes <a href=\"https:\/\/www.publicdomainpictures.net\/en\/view-image.php?image=137873&amp;picture=potato\">Potato<\/a> by Charles Rondeau, <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/\">public domain (CC0)<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:McDonalds-French-Fries-Plate.jpg\">McDonalds-French-Fries-Plate<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Evan-Amos\">Evan-Amos<\/a>, <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/\">public domain (CC0)<\/a>; <a href=\"https:\/\/pdpics.com\/photo\/2316-potato-chips-bowl\/\">Potato chips bowl<\/a> by pdpics.com, <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/\">public domain (CC0)<\/a>.]<\/caption>\n<thead>\n<tr style=\"height: 47px\">\n<td style=\"width: 20.0464%;height: 47px\"><\/td>\n<td style=\"width: 25.4644%;height: 47px\">\n<p style=\"text-align: center\"><strong>Baked Potato<\/strong><br \/>\n[based, skin on, plain]<\/p>\n<\/td>\n<td style=\"width: 27.4768%;height: 47px;text-align: center\"><strong>French Fries<\/strong><br \/>\n[Medium order]<\/td>\n<td style=\"width: 27.0124%;height: 47px\"><strong>Potato Chips<\/strong><br \/>\n[1 oz. serving of 11 chips]<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr style=\"height: 15px\">\n<td style=\"width: 20.0464%;height: 15px\">Calories<\/td>\n<td style=\"width: 25.4644%;height: 15px;text-align: center\">110<\/td>\n<td style=\"width: 27.4768%;height: 15px;text-align: center\">340<\/td>\n<td style=\"width: 27.0124%;height: 15px\"><\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"width: 20.0464%;height: 15px\">Calories from fat<\/td>\n<td style=\"width: 25.4644%;height: 15px;text-align: center\">0<\/td>\n<td style=\"width: 27.4768%;height: 15px;text-align: center\">144<\/td>\n<td style=\"width: 27.0124%;height: 15px\"><\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"width: 20.0464%;height: 15px\">Fat<\/td>\n<td style=\"width: 25.4644%;height: 15px;text-align: center\">0g<\/td>\n<td style=\"width: 27.4768%;height: 15px;text-align: center\">16g<\/td>\n<td style=\"width: 27.0124%;height: 15px\"><\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"width: 20.0464%;height: 15px\">Carbohydrates<\/td>\n<td style=\"width: 25.4644%;height: 15px;text-align: center\">26g<\/td>\n<td style=\"width: 27.4768%;height: 15px;text-align: center\">44g<\/td>\n<td style=\"width: 27.0124%;height: 15px\"><\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"width: 20.0464%;height: 15px\">Protein<\/td>\n<td style=\"width: 25.4644%;height: 15px;text-align: center\">3g<\/td>\n<td style=\"width: 27.4768%;height: 15px;text-align: center\">4g<\/td>\n<td style=\"width: 27.0124%;height: 15px\"><\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"width: 20.0464%;height: 15px\">Sodium<\/td>\n<td style=\"width: 25.4644%;height: 15px;text-align: center\">0g<\/td>\n<td style=\"width: 27.4768%;height: 15px;text-align: center\">230mg<\/td>\n<td style=\"width: 27.0124%;height: 15px\"><\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"width: 20.0464%;height: 15px\">Dietary fibre<\/td>\n<td style=\"width: 25.4644%;height: 15px;text-align: center\">2g<\/td>\n<td style=\"width: 27.4768%;height: 15px;text-align: center\">4g<\/td>\n<td style=\"width: 27.0124%;height: 15px\"><\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"width: 20.0464%;height: 15px\">Sugars<\/td>\n<td style=\"width: 25.4644%;height: 15px;text-align: center\">1g<\/td>\n<td style=\"width: 27.4768%;height: 15px;text-align: center\">0g<\/td>\n<td style=\"width: 27.0124%;height: 15px\"><\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"width: 20.0464%;height: 15px\">Cholesterol<\/td>\n<td style=\"width: 25.4644%;height: 15px;text-align: center\">0g<\/td>\n<td style=\"width: 27.4768%;height: 15px;text-align: center\">0g<\/td>\n<td style=\"width: 27.0124%;height: 15px\"><\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"width: 20.0464%;height: 15px\">Ingredients<\/td>\n<td style=\"width: 25.4644%;height: 15px;text-align: center\">Potato<\/td>\n<td style=\"width: 27.4768%;height: 15px;text-align: center\">Potatoes, vegetable oil (canola oil), soybean oil, hydrogenated soybean oil, natural beef flavour (wheat and milk derivatives), citric acid (preservative), dextrose, sodium acid pyrophosphate (main colour), salt.<\/td>\n<td style=\"width: 27.0124%;height: 15px\">Potatoes, vegetable oil (sunflower, corn, and\/or canola oil), bacon and cheddar loaded potato skins seasoning (maltodextrin - made from corn) salt, cheddar cheese (milk, cheese cultures, salt enzymes), sour cream (cultured cream, skin milk), whey, dried onion, monosodium.<\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\">Not only have we transformed the food supply and our eating in ways that are detrimental to our health, but these changes have been accompanied by reductions in physical activity. <strong>Sedentarism<\/strong> is built into contemporary lifestyles. Think of how much time you spent sitting down today. Some of it may have been in class or at work, some may have been driving a car or perhaps binge-watching your favourite show, playing a video game, or checking in on social media. An inactive lifestyle is almost dictated by the digital age (Lucock Et al<em>.<\/em> 2014). Levels of physical activity in many countries are now so low that large portions of the population are completely sedentary, including one in five Americans (CDC 2022). For a species whose biology evolved in an environment where walking, lifting, and carrying were part of daily life, this is unhealthy and often leads to weight gain.<\/p>\n<h2 class=\"import-Normal\">Biology and Genetics of Weight<\/h2>\n<figure style=\"width: 344px\" class=\"wp-caption alignright\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image5-7.png\" alt=\"Four individuals of various ages walk alongside trees.\" width=\"344\" height=\"229\" \/><figcaption class=\"wp-caption-text\">Figure 17.6: Participants of a walk against diabetes and for general fitness around Nauru airport. Credit: <a href=\"https:\/\/www.flickr.com\/photos\/106853342@N04\/10709038046\">Participants of a walk against Diabetes and for general fitness around Nauru airport<\/a> by Lorrie Graham, <a href=\"https:\/\/www.flickr.com\/photos\/dfataustralianaid\/\">Department of Foreign Affairs and Trade<\/a>, is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\">CC BY 2.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Research indicates multiple genetic variants influence weight gain, and they are not spread evenly among human populations. Tuomo Rankinen and colleagues (2006) identified 127 genes associated with obesity, of which 22 contributed to weight gain. Claude Bouchard (2007) then identified five categories of obesity-promoting genotypes. These genotypes promote sedentarism, result in low metabolism, and lead to poor regulation of appetite, and a propensity to overeat. An example of the impact such genotypes can have in an environment of plenty is found among the population of the Micronesian island of Nauru. Historically, the island was geographically isolated and the food supply was unpredictable. These conditions favoured genotypes that promoted the ability to rapidly build up and store fat in times of food availability. In Nauruans, there are two genetic variants favouring weight gain and insulin resistance, and both are associated with obesity and type 2 diabetes. One variant is also associated with hypertension. One of these variants is also found in Pima Indians, who live in parts of Arizona and Mexico. In the Pima, this variant is associated with a high <strong>b<\/strong><strong>ody mass index (BMI) <\/strong> and type 2 diabetes, although it is not associated with the same outcomes in Japanese and British subjects (de Silva Et al<em>.<\/em> 1999). The other variant was analysed in Finnish and South Indian populations, neither of whom experienced the same outcome as Nauruans. This suggests these alleles may act as modifying genes for type 2 diabetes in some population groups (Baker Et al. 1994). Unfortunately, Nauruans are one of those groups. Eventually, they became wealthy through phosphate mining on the island, gaining access to a calorie-rich Western diet of imported foods and developing a sedentary lifestyle. This resulted in rates of type 2 diabetes as high as 30\u201340% in Nauruans over the age of 15, which became the leading cause of death (Lucock Et al. 2014), something Nauruans are taking seriously (Figure 17.6).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Factors other than biology influence which populations that carry a genetic predisposition to diabetes actually express it. The Onk Akimel O\u2019odham (Pima) of Arizona, for example, were seriously impacted by U.S. government policies that affected water rights, forcing the population away from subsistence farming to dependence on government handouts and convenience food. This resulted in a significant loss of physical activity, malnutrition, and obesity. The Onk Akimel O\u2019odham continue to experience hardship due to high rates of unemployment, poverty, and depression, sometimes made worse by alcoholism. In the absence of these pressures, the Onk Akimel O\u2019odham were diabetes free for centuries prior, even though they relied on agriculture for subsistence, suggesting genetics alone is not responsible for high rates of obesity and diabetes in current Onk Akimel O\u2019odham populations (Smith-Morris 2004).<\/p>\n<figure style=\"width: 372px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image3-4-1.jpg\" alt=\"A human body outline with multiple points, each listing related medical complications.\" width=\"372\" height=\"493\" \/><figcaption class=\"wp-caption-text\">Figure 17.7: Medical complications of obesity include stroke, sleep apnea, lung disease, liver disease, gallstones, cancer, heart disease, diabetes, pancreatitis, abnormal periods and infertility, arthritis, inflamed veins, and gout. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Medical_complications_of_obesity.png\">Medical complications of obesity<\/a> by the <a href=\"https:\/\/www.cdc.gov\/\">Centers for Disease Control and Prevention (CDC)<\/a> has been modified (colour changed and cancer list shortened) 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\">Obesity also has an epigenetic component. You learned about epigenetics in Chapter 4. With regard to obesity, epigenetics is counterintuitive in that mothers who do not take in enough calories during pregnancy often give birth to babies who grow up to be fat. What takes place is the fetus receives signals during pregnancy from its mother through the placenta and intrauterine environment about environmental conditions outside of the womb, in this case food insecurity. These signals encourage the turning on and off of genes related to metabolism. This alters the phenotype of the fetus so that if the child is born into an environment where food is plentiful, it will put on weight rapidly whenever possible, leading to obesity and related diseases later in life. If the child is a girl, her own eggs are formed in utero with the same genetic changes coded in, meaning she will pass along this same genetic predisposition to gain weight to her children. Hence, a biological propensity toward obesity can continue across generations (Worthman &amp; Kuzara 2005). Epigenetic changes to genes that promote weight gain are argued to be partly responsible for the rapid rise in obesity and diabetes in developing countries gaining access to Western diets (Stearns, Nesse, and Haig 2008).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Obesity and overweight put a strain on several biological systems of the body, including the <strong>circulatory<\/strong>, endocrine, and skeletal systems, contributing to hypertension, heart disease, <strong>stroke<\/strong>, diabetes, and osteoarthritis (Figure 17.7). Obesity also elevates the risk of cancers of the breast, endometrium, kidney, colon, esophagus, stomach, pancreas, and gallbladder (National Institutes of Health 2017; Vucenik &amp; Stains 2012). Diabetes\u2014one of the fastest-growing health conditions around the globe (WHO 2016) and one tightly connected to obesity and overweight\u2014is the focus of the following Special Topics box.<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Diabetes<\/h2>\n<p class=\"import-Normal\"><strong>Diabetes mellitus<\/strong> is an endocrine disorder characterized by excessively high blood glucose levels (Martini Et al<em>.<\/em> 2013). According to a report released by the World Health Organization, the number of people living with diabetes is growing in all regions of the world. Rates of diabetes have nearly doubled in the past three decades, largely due to increases in obesity and sugary diets (WHO 2016). One in 10 people around the world, 537 million people, now have diabetes, and three out of four live in low- and middle-income countries (IDF 2022). In the United States, 37 million people have diabetes (CDC 2020), where the disease is rising fastest among millennials (those ages 20\u201340) (BCBSA 2017), and one in every two adults with diabetes is undiagnosed (IDF 2022). Obesity and diabetes are linked: obesity causes a diet-related disease (diabetes) because of humans\u2019 evolved metabolic homeostasis mechanism, which is poorly suited to contemporary energy environments (Lucock Et al<em>.<\/em> 2014).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">To function properly, cells need a steady fuel supply. Blood sugar (glucose) is the fuel for most cells in the body, and the body produces the hormone <strong>insulin<\/strong> to help move glucose into cells that need it (Figure 17.8). Foods that most readily supply glucose to your bloodstream are carbohydrates, especially starchy foods like potatoes or sweet, sugary foods like candy and soda. The body can also convert other types of foods, including protein-rich foods (e.g., lean meats) and fatty foods (e.g.<em>,<\/em> vegetable oils and butter), into blood sugar in the liver via gluconeogenesis. Insulin\u2019s main job is to tell your cells when to take up glucose. The cell also has to listen to the signal and mobilize the glucose transporters. This not only allows your cells to get the energy they need, but it also keeps blood sugar from building up to dangerously high levels when you are at rest.<\/p>\n<figure style=\"width: 485px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image7-7.png\" alt=\"The cycle of how a body with diabetes processes nutrients from food.\" width=\"485\" height=\"648\" \/><figcaption class=\"wp-caption-text\">Figure 17.8: Carbohydrates are eaten and broken down into simple sugars (e.g., glucose). Glucose enters the bloodstream from the intestines, and the increase in glucose stimulates the pancreas to release insulin into the bloodstream. Insulin deposits glucose in the muscles and fat cells, where it is stored and used for energy. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-11\/\">Glucose metabolism (Figure 16.7)<\/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\">This system has limits. Like the rest of our biology, it evolved during several million years when sugar was hard to come by and carbohydrates took the form of fresh foods with a low <strong>glycemic index (GI)<\/strong>. Our ancestors were also active throughout the day, taking pressure off of the endocrine system. Now, sedentary lifestyles and processed-food diets cause many of us to take in more calories\u2014and especially more carbohydrates\u2014than our bodies can handle. There is only so much blood sugar your cells can absorb. Many modern populations are taxing those limits. After years of being asked by insulin to take in more glucose than they can use, cells eventually stop responding (McKee &amp; McKee 2015). This is called type 2 diabetes or insulin resistance, which accounts for 90\u201395% of diabetes cases in the United States (Figure 17.9). type 1 diabetes is believed to be caused by an autoimmune response in which your immune system is attacking and destroying the insulin-producing cells in your pancreas (Figure 17.9). type 1 diabetes is a genetic condition that often shows up early in life, while type 2 is more lifestyle-related and develops over time.<\/p>\n<figure style=\"width: 462px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image9-6.png\" alt=\"Type 1 cells and Type 2 cells and their response to insulin.\" width=\"462\" height=\"276\" \/><figcaption class=\"wp-caption-text\">Figure 17.9: Type 1 and type 2 diabetes. For Type 1 Diabetes, cells do not absorb glucose becuase there is no insulin. For Type 2 Diabetes, although there is insulin available, cells do not respond to it. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-11\/\">Type 1 and Type 2 Diabetes (Figure 16.8)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson has been modified (text) and 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\">Cardiovascular Disease<\/h2>\n<p class=\"import-Normal\">Cardiovascular disease (CVD)\u2014which includes coronary heart disease, hypertension (high blood pressure), and stroke\u2014is the leading cause of death globally, and heart disease remains the number one cause of death in the United States (American Heart Association 2018). Risk factors for cardiovascular disease include diet, obesity\/overweight, diabetes, smoking and alcohol consumption, and physical inactivity.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The connections between these factors and heart disease may not seem obvious and will be addressed here beginning with diet. Diets high in saturated fat and cholesterol can lead to atherosclerosis, a condition in which fat and cholesterol form plaque inside the arteries, eventually building up and hardening to the point that blood flow is blocked. Too much salt in the diet leads to fluid retention, which increases blood volume and thereby blood pressure, taxing the heart. Obesity\/overweight contribute to cardiovascular disease directly through increases in total blood volume, cardiac output, and cardiac workload. In other words, the heart has to work much harder if one is overweight (Akil &amp; Ahmad 2011). Obesity also relates to CVD indirectly through elevation of blood pressure (hypertension) and diabetes. High levels of blood glucose from diabetes can damage blood vessels and the nerves that control the heart and blood vessels. Alcohol consumption can raise blood pressure and triglyceride levels, a type of fat found in the blood. Alcohol also adds extra calories, which may cause weight gain, especially around the abdomen, which is directly associated with risk of a heart attack (Akil &amp; Ahmad 2011). Cigarette smoking also increases the risk of coronary heart disease. Nicotine increases blood pressure; in addition, cigarette smoke causes fatty buildup in the main artery in the neck and thickens blood, making it more likely to clot. It also decreases levels of HDL (\u201cgood\u201d) cholesterol (American Heart Association 2018). Even secondhand smoke can have an adverse effect if exposure occurs on a regular basis. Chronic psychological stress also elevates the risk of heart disease (Dimsdale 2008). The repeated release of stress hormones like adrenaline elevates blood pressure and may eventually damage artery walls. The human <strong>stress response<\/strong> and its connections to health and disease are discussed in more detail below.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">However, physical activity alters the likelihood of having heart disease, both directly and indirectly. Regular exercise of moderate to vigorous intensity strengthens the heart muscle and allows capillaries, tiny blood vessels in your body, to widen, improving blood flow. Regular exercise can also lower blood pressure and cholesterol levels and manage blood sugar levels, all of which reduce the risk of CVD.<\/p>\n<h2 class=\"import-Normal\">Cancer<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Cancer is the second-leading cause of death globally, causing one in every six deaths and killing nearly nine million people in 2015 (WHO 2018). Lifetime cancer risk in developed Western populations is now one in two, or 50% (Greaves 2015). Approximately one-third of deaths from cancer are due to behavioural and dietary factors, including high body mass index (BMI), low fruit and vegetable intake, lack of physical activity, and the use of tobacco and alcohol. Depending on the type of cancer and one\u2019s own genetic inheritance, these factors can increase cancer risk from 2- to 100-fold (Greaves 2015). Cancer is the result of interactions between a person's genes and three categories of external agents: physical carcinogens (e.g., ultraviolet radiation), chemical carcinogens (e.g., tobacco smoke, asbestos), and biological carcinogens, such as infections from certain viruses, bacteria, or parasites (WHO 2018). Obesity is also a risk factor for cancer, including of the breast, endometrium, kidney, colon, esophagus, stomach, pancreas, and gallbladder (National Institutes of Health 2017; Vucenik &amp; Stains 2012).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Cancer has been regarded as a relatively recent affliction for humans that became a problem after we were exposed to modern carcinogens and lived long enough to express the disease (David &amp; Zimmerman 2010). Given the long history that humans share with many oncogenic (cancer-causing) parasites and viruses (Ewald 2018), and the recent discovery of cancer in the metatarsal bone of a 1.8-million-year-old hominin (Odes Et al. 2016), this view is being challenged (See \u201cSpecial Topic: Life Choices and Reproductive Cancers in Women\u201d). The difficulties of identifying cancer in archaeological populations are many. Most cancer occurs in soft tissue, which rarely preserves, and fast-growing cancers would likely kill victims before leaving evidence in bone. It is also difficult to distinguish cancer from benign growths and inflammatory disease in ancient fossils, and there is often postmortem damage to fossil evidence from scavenging and erosion. However, using 3-D images, South African researchers recently diagnosed a type of cancer called osteosarcoma in a toe bone belonging to a human relative who died in Swartkrans Cave between 1.6 and 1.8 million years ago (Randolph-Quinney Et al. 2016). This study provides the earliest evidence of cancer in hominins.<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Life Choices and Reproductive Cancers in Women<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Behavioural or \u201clifestyle\u201d choices have an impact on cancer risk. Breast cancer is one example. It is the most common cancer in women worldwide, but <strong>incidence<\/strong> of new cases varies from 19.3 per 100,000 women in Eastern Africa to 89.7 per 100,000 women in Western Europe (WHO 2018). These differences are attributable to cultural changes among women in Western, industrialized countries that are a mismatch for our evolved reproductive biology. Age at <strong>menarche<\/strong>, the onset of menstrual periods, has dropped over the course of the last century from 16 to 12 years of age in the U.S. and Europe, with some girls getting their periods and developing breasts as young as eight years old (Greenspan &amp; Deardorff 2014, Figure 17.10). A World Health Organization study involving 34 countries in Europe and North America suggests the primary reason for the increase in earlier puberty is obesity, with differences in BMI accounting for 40% of individual- and country-level variance (Currie Et al<em>.<\/em> 2012). Early puberty in girls is associated with increased risk of breast cancer, ovarian cancer, diabetes, and high cholesterol in later life (Pierce &amp; Hardy 2012).<\/p>\n<figure style=\"width: 554px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image12-3.jpg\" alt=\"A graph shows the decrease in age at menarche for five European nations and United States.\" width=\"554\" height=\"434\" \/><figcaption class=\"wp-caption-text\">Figure 17.10: Decreasing ages at time of first menstruation in selected countries. Credit: <a href=\"https:\/\/en.wikipedia.org\/wiki\/Menarche#\/media\/File:Acceleration1.jpg\">Acceleration1.jpg<\/a> by Yahadzija 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>&nbsp;<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">At the same time that age at puberty is dropping for girls in Western nations, age at birth of the first child is later, at 26 years old (Mathews and Hamilton 2016). Women are also having fewer children, two on average (Gao 2015), with 15% of women choosing to remain childless (Livingston 2015). Rates of breastfeeding have risen in recent decades but drop to only 27% of infants once babies reach 12 months of age (CDC 2014). In contrast, data from modern foraging populations (Eaton Et al. 1994) indicate that age at menarche is around 16 years old, age at birth of the first child is 19, breastfeeding on demand continues for three years for each child, and the number of children averages six. These differences relate to elevated risk for reproductive cancers, including breast cancer, among women in developed countries.<\/p>\n<p class=\"import-Normal\">Other than an established genetic risk (e.g., BRCA gene), the primary risk factor for breast cancer is exposure to estrogen. For women living in modern, industrialized economies, this exposure now often comes from women\u2019s own ovaries rather than from external environmental sources (Stearns, Nesse, and Haig 2008). Women in cultures without contraception are pregnant or breastfeeding for much of their reproductive lives, resulting in 100 or so menstrual cycles per lifetime. In contrast, Western women typically experience 400 or more (Strassmann 1997). This is partly due to early puberty. From menarche to the birth of a woman\u2019s first child can be 14 years or longer in Western populations, after which breastfeeding, if undertaken at all, lasts for a few weeks or months. Oral contraceptives or other hormonal methods to control reproduction induce monthly periods. Age at menopause (the cessation of menstrual cycles) is 50\u201355 years old across human populations. For Western women, this translates into forty years of menstrual cycling. Each month, the body prepares for a pregnancy that never occurs, experiencing cell divisions that put women at risk for cancers of the breast, endometrium, ovaries, and uterus (Strassmann 1999). Obesity adds to the risk, as adipose (fat) tissues are the primary source of estrogen biosynthesis. Thus, weight gain during the postmenopausal stage means higher exposure to estrogen and a greater risk of cancer (Ali 2014).<\/p>\n<p class=\"import-Normal\">Women cannot return to our evolutionary past, and there are significant social and economic reasons for delaying pregnancy and having fewer children. These include achieving educational and career goals, greater earning power, a reduction in the gender pay gap, more enduring marriages, and a decrease in the number of women needing public assistance (Sonfield Et al. 2013). There are also cultural means by which we might reduce the risk of reproductive cancers. These include reformulating hormonal contraceptives with enough estrogen to maintain bone density but reducing the number of menstrual periods over the reproductive lifespan (Stearns, Nesse, &amp; Haig 2008). Reducing fat intake may also lower estrogen levels. High-fat diets contribute to breast tumor development, while high fibre diets are beneficial in decreasing intestinal resorption of estrogenic hormones. Exercise also appears protective. Studies of former college athletes demonstrate risks of breast, uterine, and ovarian cancers later in life, two to five times lower than those of nonathletes (Eaton Et al<em>.<\/em> 1994).<\/p>\n<\/div>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Stress<\/h2>\n<p class=\"import-Normal\">Have you ever been \u201cstressed out\u201d in class? Say you\u2019re in a large lecture hall with a hundred other people, or even in a small class where you don\u2019t know anyone. You\u2019re not sure about something the professor just said and you would really like to ask about it, so you start to raise your hand. Does your heart begin to pound and your mouth become dry? Do you get so nervous that you choose to ask a classmate after lecture instead? If so, you are not alone. Fear of speaking in public is one of the most common social phobias (APA 2013). It has been estimated that 75% of all people experience some degree of anxiety or nervousness when it comes to public speaking (Hamilton 2011), and surveys have shown that most people fear public speaking more than they fear death (Croston 2012).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">We have evolution to thank for this.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Humans, like other primates, are social animals. Being part of a group helped us to survive predation, get enough to eat, and successfully raise our young. When faced with standing up in front of a group, or even speaking up in class, we break into a sweat because we are afraid of rejection. Psychologist Glenn Croston (2012) writes, \u201cThe fear is so great because we are not merely afraid of being embarrassed or judged. We are afraid of being rejected from the social group, ostracized and left to defend ourselves all on our own. We fear ostracism still so much today it seems, fearing it more than death, because not so long ago getting kicked out of the group probably really was a death sentence.\u201d Hence, it is no surprise that public speaking triggers a stress response among much of humankind.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The human nervous system evolved in a context where quick responses to perceived threats presented an evolutionary advantage. The \u201cfight or flight\u201d response was honed during millions of years when threats more often took the form of an approaching lion than an approaching deadline. Our body\u2019s stress response, however, is triggered by a wide variety of stressors that produce the same general pattern of hormonal and physiological adjustments (Martini Et al. 2013). In today\u2019s world, the system is often stuck in the \u201con\u201d position due to the constant pressures of modern life, and this is a significant influence on health and disease.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">It is important to recognize that there are different types of stress and the time in life when adult coping mechanisms are formed is in childhood. In children, some stressors can be positive\u2014for example, stressors that are mild to moderate in magnitude, and accompanied by the support of a caring adult, which help children develop pathways by which stress is dealt with by the body throughout life. In a young child, a positive stress response might be going to the pediatrician to receive a vaccination and receiving encouragement and comfort from both parent and practitioner. A tolerable stress response is more serious, precipitated by something like a divorce or death of a relative. Again, buffered by positive support from surrounding adults, these types of stressors can be successfully managed by children. Toxic stress, however, \u201cresults from strong, frequent or prolonged activation of the stress response in the absence of the buffering protection of a supportive adult relationship\u201d (Shonkoff &amp; Garner 2012). Examples include child abuse or neglect, parental substance abuse, homelessness, and violence. In the absence of adequate psychological and physical support, the biological pathways of a child\u2019s physiological stress response are altered and lead to reduced abilities to cope with life\u2019s challenges as an adult.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The negative effects of sustained, elevated cortisol levels on health are well documented. These include higher levels of infectious disease and slowed growth in childhood (Flinn &amp; England 2003) and increased incidence of heart disease, obesity, and diabetes in adults (Worthman &amp; Kuzara 2005). Contrary to our evolutionary past, many causes of sustained stress in contemporary societies are psychosocial rather than physical threats. These can include an unhappy marriage or frustrations at work (Dimsdale 2008). Stressors can also be more subtle. For example, a review of research into the effects of stress on health indicated that experiencing racism was a significant stressor that was associated with alcohol consumption, psychological distress, overweight, abdominal obesity, and higher fasting-glucose levels among minority groups (Williams &amp; Mohammed 2013). Chronic, everyday racial discrimination is also associated with the hardening of coronary arteries, elevated blood pressure, giving birth to lower-birth-weight infants, cognitive impairment, poor sleep, and visceral fat, which is fat stored deep inside the belly, wrapped around the organs, including the liver and intestines. Visceral fat is a sign of <strong>m<\/strong><strong>etabolic syndrome<\/strong>, increasing the risk of stroke, heart disease, and type 2 diabetes. These effects have been shown to increase morbidity and mortality among members of affected groups.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Epigenetics can also be a factor in how a person is able to deal with stressful situations. Maternal experiences of stress during pregnancy have the potential to permanently alter the physiology of mothers\u2019 offspring, especially the hypothalamic-pituitary-adrenal (HPA) axis. The HPA axis regulates metabolism, blood pressure, and the immune response, and these alterations can predispose prenatally stressed individuals to suffer metabolic, cardiovascular, and mental disorders in adulthood (Palma-Gudiel Et al. 2015). These experiences carry across generations, with children of Holocaust survivors who experienced PTSD demonstrating similar changes in neurochemistry in the absence of a sustained, traumatic event, as did infant offspring of mothers who developed PTSD during pregnancy after witnessing the traumatic events of 9\/11 (Yehuda &amp; LeDoux 2007).<\/p>\n<h2 class=\"import-Normal\" style=\"text-indent: 0pt\">Syndemics and the Ecological Model<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">It is important to recognize that disease risk is not spread evenly within or between populations. Diseases combine and interact to create a <strong>syndemic<\/strong>, where the coexistence of two or more conditions exacerbates the effects of one or all conditions. A syndemic (versus a pandemic, for example) takes into account social, political, economic, and environmental factors that increase risk for the clustering of two or more diseases (Singer Et al. 2017). One of the first syndemics identified involved substance abuse, violence, and AIDS. In inner cities in the U.S., the health crisis around HIV\/AIDS was related to tuberculosis, sexually transmitted infections, hepatitis, cirrhosis, infant mortality, drug abuse, suicide, and homicide. These were connected to poverty, homelessness, unemployment, poor nutrition, lack of social support, and social and ethnic inequality (Singer Et al. 2017). Together, these factors and others, like health policy and unequal access to health care, form an <strong>ecological model<\/strong> of health and disease, one that moves beyond biology and medical intervention (Sallis Et al. 2008).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The COVID-19 pandemic represents a syndemic in which systemic racism in the healthcare system, differential access to diagnosis and treatment, income, employment, housing, family structure, pre existing conditions, and public health policies combined to result in higher rates of infection and death for African Americans, Native Americans, Asians, and Hispanic populations in the United States (Figure 17.11).<\/p>\n<table class=\"grid\" style=\"border-collapse: collapse;width: 100%\" border=\"0\">\n<caption>Figure 17.11: Risk for COVID-19 infection, hospitalization, and death by race\/ethnicity. Race and ethnicity are risk markers for other underlying conditions that affect health, including socioeconomic status, access to health care, and exposure to the virus related to occupation, e.g., frontline, essential, and critical infrastructure workers. Credit: <a href=\"https:\/\/www.cdc.gov\/coronavirus\/2019-ncov\/covid-data\/investigations-discovery\/hospitalization-death-by-race-ethnicity.html\">Risk for COVID-19 Infection, Hospitalization, and Death by Race\/Ethnicity<\/a> by the <a href=\"https:\/\/www.cdc.gov\/\">Centers for Disease Control and Prevention<\/a> is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/caption>\n<thead>\n<tr class=\"shaded\">\n<td style=\"width: 130.367px\">Rate ratios compared to White, Non-Hispanic persons<\/td>\n<td style=\"width: 130.367px\">American Indian or Alaska Native, Non-Hispanic persons<\/td>\n<td style=\"width: 130.367px\">Asian, Non-Hispanic persons<\/td>\n<td style=\"width: 130.383px\">Black or African American, Non-Hispanic persons<\/td>\n<td style=\"width: 130.35px\">Hispanic or Latino persons<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr>\n<td style=\"width: 130.367px\">Cases<\/td>\n<td style=\"width: 130.367px\">1.6x<\/td>\n<td style=\"width: 130.367px\">.8x<\/td>\n<td style=\"width: 130.383px\">1.1x<\/td>\n<td style=\"width: 130.35px\">1.5x<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 130.367px\">Hospitalization<\/td>\n<td style=\"width: 130.367px\">2.7x<\/td>\n<td style=\"width: 130.367px\">.8x<\/td>\n<td style=\"width: 130.383px\">2.3x<\/td>\n<td style=\"width: 130.35px\">2.0x<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 130.367px\">Death<\/td>\n<td style=\"width: 130.367px\">2.1x<\/td>\n<td style=\"width: 130.367px\">.8x<\/td>\n<td style=\"width: 130.383px\">1.7x<\/td>\n<td style=\"width: 130.35px\">1.8x<\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<p class=\"import-Normal\">COVID-19 was the third leading cause of death in the U.S. in 2020 and 2021 (NIH 2022; Figure 17.12), but morbidity and mortality was not equally spread across the population. Working-class people and people of colour in the U.S. are more likely to live in poverty, in areas with high rates of crime and violence, and in close proximity to freeways and environmental threats like petrochemical plants and waste incinerators (Singer and Baer 2012). Many such neighbourhoods are also food \u201cdeserts\u201d without ready access to a healthy, affordable diet, made more challenging by residents not owning a car (Food Empowerment Project n.d.). Low-income people also often lack access to high-quality health care and delay or avoid preventive care and health screenings (Ross Et al. 2007). These factors contributed to higher rates of preexisting conditions, including obesity, diabetes, hypertension, asthma, heart disease, chronic obstructive pulmonary disease (COPD), and smoking behaviour, which then led to more complications and higher death rates from COVID (Ghosh Et al. 2021).<\/p>\n<p class=\"import-Normal\">Family structure also affected COVID exposure and severity. Many Americans live in multigenerational households, including 27% of Hispanics, 29% of Asians, 26% of African Americans, and 20% of Whites (Cohn &amp; Passel 2018). Not all multigenerational households are equal, however. Over twice as many African Americans as Whites are in multigenerational families in which at least one family member is unemployed, and over three times as many African Americans are in multigenerational families in which everyone is simultaneously unemployed (Park, Wiemers, and Seltzer 2019). Family members in multigenerational households were at a much higher risk of developing more severe forms of COVID due to decreased personal space and multiple exposures to the virus, as well as higher rates of diabetes, smoking, and residents living below the poverty line (Ghosh Et al. 2021). While aimed at reducing overall infection rates from COVID, public health measures such as mandatory lockdowns only exacerbated the situation in overcrowded and multigenerational housing, resulting in higher rates of infection and death in these communities.<\/p>\n<div style=\"margin: auto\">\n<table class=\" aligncenter\" style=\"width: 468pt;height: 195px\">\n<caption>Figure 17.12: Top five causes of death in the U.S. and worldwide since 2020. Credit: Top five causes of death in the U.S. and worldwide original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Joylin Namie is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Based on data from Shiels Et al. 2022 and Traeger 2022.<\/caption>\n<thead>\n<tr class=\"shaded\" style=\"height: 30px\">\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 164.6px\">\n<p class=\"import-Normal\"><strong>United <\/strong><strong>States<\/strong><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 429.733px\">\n<p class=\"import-Normal\"><strong>Worldwide<\/strong><\/p>\n<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table2-R\" style=\"height: 0\">\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 164.6px\">\n<p class=\"import-Normal\">1. Heart disease<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 429.733px\">\n<p class=\"import-Normal\">1. Heart disease<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 0\">\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 164.6px\">\n<p class=\"import-Normal\">2. Cancer<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 429.733px\">\n<p class=\"import-Normal\">2. Stroke<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 0\">\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 164.6px\">\n<p class=\"import-Normal\">3. COVID-19<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 429.733px\">\n<p class=\"import-Normal\">3. COVID-19<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 0\">\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 164.6px\">\n<p class=\"import-Normal\">4. Accidents<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 429.733px\">\n<p class=\"import-Normal\">4. Chronic Obstructive Pulmonary Disease<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 0\">\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 164.6px\">\n<p class=\"import-Normal\">5. Stroke<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 429.733px\">\n<p class=\"import-Normal\">5. Lower respiratory infections<\/p>\n<\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"height: 15px;width: 165.467px\"><\/td>\n<td style=\"height: 15px;width: 430.6px\"><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<p class=\"import-Normal\">There is a long history of systemic racism and discrimination in the medical system in the United States (Washington 2006). African Americans have been subjected to medical testing and experimentation without their consent or knowledge since the time of slavery. They continue to routinely receive care of poorer quality than whites (Williams &amp; Wyatt 2015), less pain medication during treatment and hospitalization (Green Et al. 2003), and differential treatment during pregnancy and childbirth (Washington 2006). Many Americans, including 50% of White medical students and residents in one recent study (Hoffman Et al. 2016), hold at least one false belief about African Americans, including \u201cBlack people\u2019s skin is thicker than white people\u2019s skin,\u201d \u201cBlacks have stronger immune systems than whites,\u201d and \u201cBlacks\u2019 nerve endings are less sensitive than whites\u2019.\u201d Such beliefs affect health care for African Americans in medical emergencies and for chronic conditions.<\/p>\n<p class=\"import-Normal\">During the COVID-19 pandemic, patients with darker skin in the United States were negatively affected by the very medical device most commonly used to assess oxygen levels in their blood. The pulse oximeter, a small device that clips onto the tip of your index finger and measures blood oxygen levels, experienced increased use in home, clinical, and hospital settings during the COVID-19 pandemic. Decisions regarding treatment and hospital admission for patients infected with COVID were often based on pulse oximeter readings (Valbuena, Merchant, &amp; Hough 2022). The problem is the device overestimates oxygen saturation in patients with darker skin, an issue which has been recognized for over thirty years (Valbuena, Merchant, &amp; Hough 2022). It would be as if a standard thermometer reported lower body temperatures for patients of colour, making it seem as if they did not have a fever when they actually did. In the case of COVID-19, Asians, Hispanics, and African Americans experienced inaccurately high readings of their oxygen levels (with African Americans and darker-skinned Hispanics having the highest), resulting in delays in treatment, hospital admission, and access to medications to treat COVID and contributing to higher severity of illness and higher death rates among these populations in comparison to whites (Fawzi Et al. 2022).<\/p>\n<p class=\"import-Normal\">Employment was also a factor in unequal exposure to and death from COVID-19 (Raifman, Skinner, &amp; Sojourner 2022), with many low-income workers making the choice (which, realistically, may not be a choice at all) to expose themselves to COVID in order to earn the funds necessary to purchase food, housing, and other necessities. Many such workers were then forced to miss work due to COVID infection. With only 35% of low-wage workers (as opposed to 95% of high-wage workers) having paid sick leave, this left many families struggling financially. Three years into the pandemic, low-wage workers continue to have the least access to COVID vaccines and boosters. The U.S. also lacks federal workplace-safety regulations with regard to vaccine and masking mandates that other nations enforce in times of high transmission, and it does not provide high-quality masks to its essential workers. Many occupations deemed essential by the CDC during the height of the pandemic\u2014such as health care, emergency services, meat packing, agricultural work, teaching, and jobs in the hospitality sector\u2014experienced higher rates of morbidity and mortality from COVID. Many of these fields disproportionately employ people of colour (McKinsey &amp; Company 2021). Given this, future policies that address the pandemic at a structural level\u2014for example, providing monetary assistance to people who work in environments with a high risk of infection, such as cleaning, nursing, transportation, retail, restaurant work, and factory work, so that they can remain at home\u2014may function more effectively to prevent transmission and curb future outbreaks (Arnot Et al. 2020).<\/p>\n<h2 class=\"import-Normal\">Food for Thought<\/h2>\n<p class=\"import-Normal\">This chapter focused primarily on health conditions prevalent in contemporary, industrialized societies that are due, in part, to the mismatch between our evolved biology and modern environments. These are the built environment and the social environment, which together form the obesogenic environment in which unhealthy behaviours are encouraged. This chapter will close by examining each of these in a college context.<\/p>\n<figure style=\"width: 275px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image2-3.jpg\" alt=\"Four individuals in a park.\" width=\"275\" height=\"183\" \/><figcaption class=\"wp-caption-text\">Figure 17.13: Students walking around a campus. Credit: <a href=\"https:\/\/www.maxpixel.net\/Row-Four-Man-Woman-People-Walking-Together-3755342\">Row four man woman people walking together 3755342<\/a> by <a href=\"https:\/\/www.maxpixel.net\/\">MaxPixel<\/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>Consider your campus from an evolutionary perspective. To what degree does the built environment lend itself to physical activity as part of daily life? Is your campus constructed in ways that promote driving at the expense of walking or biking? If driving is necessary, is parking available close to the buildings or do you need to walk a fair distance from the parking lot to your destination? Do the buildings have stairs or ramps or is it necessary to take the elevator? Is it possible to negotiate safely around campus on foot or by bike in all weather? After dark? How about the classrooms and computer labs? Do they have standing or treadmill desks? Does your class schedule encourage walking from building to building between classes, or are most courses in your major scheduled in the same location? Most college majors also lack a physical education requirement, leaving it up to students to incorporate exercise into already-challenging schedules (Figure 17.13).<\/p>\n<p class=\"import-Normal\">Sociocultural factors that contribute to obesity include food advertising, ubiquitous fast-food and junk food options, and social pressure to consume, all of which are present on college campuses. Although nutrition on campuses has improved in recent years, many students find eating healthy in the dining halls and dorms challenging (Plotnikoff Et al. 2015), and students who live off campus fare even worse (Small Et al<em>.<\/em> 2013). There are also parties and other social events, a normal part of college life, that involve unhealthy food and encourage behaviours like alcohol consumption and smoking. Give some thought to the social atmosphere on your campus and the ways it may contribute to obesity. My own freshman orientation involved a succession of pizza parties, ice cream socials, and barbecues, followed by late-night runs to the nearest fast-food outlet. The purpose of these events was to encourage people to make friends and feel comfortable living away from home, but the foods served were unhealthy, and there was social pressure to join in and be part of the group. Such activities set students up for the \u201cfreshman fifteen\u201d and then some. They also reinforce the idea that being social involves eating (and sometimes drinking and\/or smoking).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Sedentarism and inactivity are also built into the academics of college life. Digital technology is a significant contributor to obesity. Students use laptops and cell phones to take notes, complete their work outside of class, and access social media. There are also video games, virtual reality headsets, and streaming television and movies for entertainment. The built environment of college already necessitates that students sit in class for hours each day, then sit at computers to complete work outside of class. The social environment enabled by digital technology encourages sitting around for entertainment. It is telling that we call it \u201cbinge watching\u201d when we spend hours watching our favourite shows. Doing so often involves eating, as well as multiple exposures to food advertising embedded in the shows themselves. In these ways, college contributes to the development of obesity-causing behaviours that can have negative health ramifications long after college is over (Small Et al. 2013).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In the U.S., the greatest increase in obesity is among young adults aged 18\u201329 years, a significant percentage of whom are college students (Plotnikoff Et al. 2015). Analyses of college students\u2019 behaviour across semesters shows consumption of fruits and vegetables drops over time, as does the amount of physical activity, while consumption of sugar-sweetened beverages and fast-food goes up, leading to weight gain at nearly six times the rate of the general public (Small Et al. 2013). In response, many colleges and universities have instituted programs to encourage healthier eating and more physical activity among students (Plotnikoff Et al<em>.<\/em> 2015). It is important to emphasize that neither changes in diet or exercise are effective on their own.. A 2022 study of over 340,000 British participants demonstrated that physical activity and diet quality did not individually have an impact on cardiovascular disease or cancers (Ding Et al. 2022). That is, hitting the gym won\u2019t counteract the consequences of consuming high-calorie, fatty foods, and eating kale all day can\u2019t cancel out sedentary habits.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Just as no one fad diet is going to prove healthier than another, no one type of exercise is better than another. Anything that raises your heart rate and that you enjoy doing for at least an hour each day will work. Take advantage of opportunities to build exercise into everyday life. Take the stairs, park as far away from buildings as possible, ride a bike or walk instead of driving, and take walks between classes instead of sitting down and checking your phone. As far as diets go, eating a few less unhealthy calories each day, one less soda, no sugar in your coffee, or letting that last slice of pizza go to someone else, make a difference in the long run. Little changes add up to bigger ones. We cannot change our biology, but we can certainly change our habits.<\/p>\n<div class=\"textbox shaded\">\n<h2>Summary<\/h2>\n<p data-start=\"134\" data-end=\"760\">The health problems faced by modern humans can often be understood through the lens of evolution. As early <em>Homo sapiens<\/em> evolved in environments with a wide diversity of edible plants and animals, we developed complex nutritional requirements. However, with dramatic changes in our environments and daily lives, modern humans now encounter a variety of diet-related challenges. These include shifts in dentition and intestinal morphology, changes in food production and preparation, and altered patterns of physical activity. Together, these factors have contributed to changes in the leading causes of morbidity and mortality.<\/p>\n<p data-start=\"762\" data-end=\"1565\">These health consequences are often framed through the concept of epidemiological transitions, which are influenced by diet, physical activity, population density, and exposure to zoonotic diseases linked to agriculture. It is proposed that the first transition occurred when humans shifted from foraging to food production, leading to increased exposure to new diseases and dietary changes. The second transition followed the Industrial Revolution, when improvements in lifestyle and living conditions reduced infectious diseases but led to a rise in non-communicable diseases. The third and fourth transitions, still unfolding today, are marked by the spread of drug-resistant pathogens, the re-emergence of infectious diseases, and new health challenges driven by globalization, urbanization, and climate change.<\/p>\n<p data-start=\"1567\" data-end=\"1807\">Modern health issues such as obesity, cardiovascular disease, cancer, stress-related conditions, and syndemics illustrate the ongoing interaction between our evolved biology and the cultural and physical environments in which we live.<\/p>\n<h2 class=\"import-Normal\">Review Questions<\/h2>\n<ul>\n<li class=\"import-Normal\">Why do humans like foods that are \u201cbad\u201d for them? Describe the evolutionary underpinnings of our tastes for sugar, salt, and fat.<\/li>\n<li class=\"import-Normal\">How might understanding contemporary disease from an evolutionary perspective benefit medical practitioners in treating their patients?<\/li>\n<li class=\"import-Normal\">Several risk factors for conditions like heart disease, diabetes, and cancer are referred to as \u201clifestyle factors,\u201d implying these are behavioural choices people make that put them at risk. These include unhealthy eating, lack of physical activity, smoking, and alcohol consumption. To what degree is unhealthy behaviour structured by the physical and social environment? For example, how does being a college student influence your eating habits, physical activity patterns, smoking, and consumption of alcohol?<\/li>\n<li class=\"import-Normal\">Who benefits from the global obesity epidemic? Think about how the following industries and institutions might profit from it: The medical establishment? The fitness industry? The diet industry? Fashion? Pharmaceutical companies? Food manufacturers? Advertisers?<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<h2 class=\"__UNKNOWN__\">Key Terms<\/h2>\n<div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\"><strong>Body mass index (BMI)<\/strong>: A person\u2019s weight in kilograms divided by the square of their height in metres. This is the most widely used measure for identifying obesity. The formula using kilograms and metres (or centimetres) is: weight (kg) \/ [height (m)]<sup>2 <\/sup>. The formula using pounds and inches is: 703 x weight (lbs) \/ [height (in)]<sup>2 <\/sup>. Use of the BMI is controversial for several reasons, including that it does not take into account age, bone structure, muscle mass, fat distribution, or ethnic and racial differences in body type.<\/p>\n<p class=\"import-Normal\"><strong>Cancer<\/strong>: A collection of related diseases in which some of the body\u2019s cells begin to divide without stopping and spread into surrounding tissues.<\/p>\n<p class=\"import-Normal\"><strong>Cardiovascular disease (CVD)<\/strong>: A disease of the heart and blood vessels, often related to atherosclerosis. CVD is a condition in which a substance called plaque builds up in the walls of the arteries, the blood vessels that carry blood away from the heart, which compromises the flow of blood to the heart or brain.<\/p>\n<p class=\"import-Normal\"><strong>Central nervous system<\/strong>: The complex of nerve tissues stemming from the brain and spinal cord that controls the activities of the body.<\/p>\n<p class=\"import-Normal\"><strong>Circulatory (system)<\/strong>: The biological system that circulates blood around the body via the heart, arteries, and veins, delivering oxygen and nutrients to organs and cells and carrying waste products away.<\/p>\n<p class=\"import-Normal\"><strong>Diabetes mellitus<\/strong>: An endocrine disorder in which high glucose (blood sugar) levels occur over a prolonged period of time. Blood glucose is your body\u2019s main source of energy and comes from the food you eat. Insulin, a hormone made by the pancreas, helps glucose from food get into your cells to be used for energy. Sometimes your body does not make enough\u2014or any\u2014insulin (type 1 diabetes) or does not take up insulin well (type 2 diabetes). Glucose then stays in your blood and does not reach your cells.<\/p>\n<p class=\"import-Normal\"><strong>\u201cDouble burden\u201d<\/strong>: Refers to parts of the world in which there is a prevalence of chronic disease (e.g., cancer, heart disease) while, at the same time, there are also high rates of infectious disease due to poverty, malnutrition, poor sanitation, and lack of access to health care, often accompanied by high rates of maternal and child mortality.<\/p>\n<p class=\"import-Normal\"><strong>Ecological model<\/strong>: Ecological models of health and disease emphasize environmental and policy contexts of behaviour, while incorporating social and psychological influences, rather than focusing on individual behaviours. These models encompass multiple levels of influence and can lend themselves to more comprehensive health interventions.<\/p>\n<p class=\"import-Normal\"><strong>Emerging infectious diseases (EIDs)<\/strong>: Infections that have recently appeared within a population or those whose incidence or geographic range is rapidly increasing or threatens to increase in the near future. Examples include Covid-19, Ebola, Zika, SARS, and avian (bird) flu.<\/p>\n<p class=\"import-Normal\"><strong>Endocrine system<\/strong>: Those organs in the body whose primary function is the production of hormones.<\/p>\n<p class=\"import-Normal\"><strong>Epidemiological transition<\/strong>: A transformation in patterns of disease (morbidity) and death (mortality) among a population.<\/p>\n<p class=\"import-Normal\"><strong>Glycemic index (GI)<\/strong>: A system that ranks foods on a scale from 1 to 100 based on their effect on blood-sugar levels. Carbohydrates with a low GI value (55 or less) are more slowly digested and metabolized causing a lower, slower rise in blood glucose and insulin levels.<\/p>\n<p class=\"import-Normal\"><strong>Hypertension<\/strong>: High blood pressure. Blood pressure is the force exerted by the blood against the walls of the blood vessels. In a blood pressure reading, the top number (usually higher) refers to the systolic pressure, the amount of pressure in your arteries during the contraction of your heart muscle when your heart beats. The bottom number is the diastolic pressure when your heart muscle is resting between beats. Hypertension can lead to severe health complications and increases the risk of heart attack and stroke.<\/p>\n<p class=\"import-Normal\"><strong>Incidence<\/strong>: The rate at which new cases of a disease occur in a population over a given period of time.<\/p>\n<p class=\"import-Normal\"><strong>Insulin<\/strong>: A hormone produced in the pancreas that regulates the amount of glucose in the blood. Lack of insulin or the inability to absorb insulin causes diabetes.<\/p>\n<p class=\"import-Normal\"><strong>Metabolic syndrome<\/strong>: A cluster of conditions, including increased blood pressure, high blood sugar, excess body fat around the waist, and abnormal cholesterol levels that occur together, increasing the risk of heart disease, stroke, and diabetes. Lifestyle changes like losing weight, exercising regularly, and making dietary changes can help prevent or reverse metabolic syndrome.<\/p>\n<p class=\"import-Normal\"><strong>Menarche<\/strong>: The first occurrence of menstruation.<\/p>\n<p class=\"import-Normal\"><strong>Morbidity<\/strong>: The number of cases of disease per unit of population occurring over a unit of time.<\/p>\n<p class=\"import-Normal\"><strong>Mortality<\/strong>: The number of deaths attributable to a particular cause per unit of population over a unit of time.<\/p>\n<p class=\"import-Normal\"><strong>Noncommunicable diseases (NCDs)<\/strong>: Also known as chronic diseases, NCDs tend to be of long duration and are the result of a combination of genetic, physiological, environmental, and behaviour factors. The main types of NCDs are cardiovascular diseases (like heart attacks and stroke<strong>)<\/strong>, cancers, chronic respiratory diseases (such as chronic obstructive pulmonary disease and asthma), and diabetes.<\/p>\n<p class=\"import-Normal\"><strong>Obesity<\/strong>: A medical condition in which excess body fat has accumulated to the point that it has adverse effects on health. Although controversial due to its lack of ethnic and racial specificity, the most widely used measure for identifying obesity is the body mass index (BMI), a person\u2019s weight in kilograms divided by the square of their height in metres. A measure of 30 kg\/m<sup>2<\/sup> is considered obese and 25\u201329 kg\/m<sup>2<\/sup> is considered overweight. Distribution of body fat also matters. Fat in the abdominal region has a stronger association with type 2 diabetes and cardiovascular disease, meaning waist-to-hip ratio and waist circumference are also important indicators of obesity-related health risk.<\/p>\n<p class=\"import-Normal\"><strong>Obesogenic<\/strong>: Promoting excessive weight gain.<\/p>\n<p class=\"import-Normal\"><strong>Omnivorous<\/strong>: Able to eat and digest foods of both plant and animal origins.<\/p>\n<p class=\"import-Normal\"><strong>Osteoarthritis<\/strong>: Refers to the degeneration of joint cartilage and underlying bone, causing pain and stiffness. In the absence of previous injury, it is most common in modern populations from middle age onward.<\/p>\n<p class=\"import-Normal\"><strong>Prevalence<\/strong>: The proportion of individuals in a population who have a particular disease or condition at a given point in time.<\/p>\n<p class=\"import-Normal\"><strong>Sedentarism<\/strong>: A way of life characterized by much sitting and little physical activity.<\/p>\n<p class=\"import-Normal\"><strong>Sedentism<\/strong>: Living in groups settled permanently in one place.<\/p>\n<p class=\"import-Normal\"><strong>Stress response<\/strong>: A predictable response to any significant threat to homeostasis. The human stress response involves the <strong>Central Nervous System<\/strong> and the endocrine system acting together. Sudden and severe stress incites the \u201cflight or flight\u201d response from the autonomic nervous system in conjunction with hormones secreted by the adrenal and pituitary glands, increasing our heart rate and breathing and releasing glucose from the liver for quick energy.<\/p>\n<p class=\"import-Normal\"><strong>Stroke<\/strong>: A stroke occurs when a blood vessel leading to the brain is blocked or bursts, preventing that part of the brain from receiving blood and oxygen, leading to cell death.<\/p>\n<p class=\"import-Normal\"><strong>Syndemic<\/strong>: The aggregation (grouping together) of two or more diseases or health conditions in a population in which there is some level of harmful biological or behavioural interface that exacerbates the negative health effects of any or all of the diseases involved. Syndemics involve the adverse interaction of diseases of all types, including infections, chronic noncommunicable diseases, mental health problems, behavioural conditions, toxic exposure, and malnutrition.<\/p>\n<p class=\"import-Normal\"><strong>Tricep skinfold measurement<\/strong>: The triceps skinfold site is a common location used for the assessment of body fat using skinfold calipers. A section of skin on the posterior (back) surface of the arm that lays atop the tricep muscle is pinched between calipers. The resulting measurement is matched against a chart standardized for age and gender.<\/p>\n<p class=\"import-Normal\"><strong>\u201cTriple burden\u201d<\/strong>: A fourth epidemiological transition currently underway in which some parts of the globe are suffering from the \u201cdouble burden\u201d of infectious and chronic diseases combined with injuries and diseases related to intensifying globalization, urbanization, deforestation, and climate change.<\/p>\n<p class=\"import-Normal\"><strong>Vector-borne diseases<\/strong>: Human illnesses caused by parasites, viruses, and bacteria that are transmitted by mosquitoes, flies, ticks, mites, snails, and lice.<\/p>\n<p class=\"import-Normal\"><strong>Zoonoses<\/strong>: Diseases that can be transmitted from animals to humans.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration<strong><br \/>\n<\/strong><\/h2>\n<p>Lents, Nathan H. 2018. <em>Human Errors: A Panorama of Our Glitches, from Pointless Bones to <\/em><em>Broken Genes<\/em>. Boston: Houghton Mifflin Harcourt.<\/p>\n<p>Stearns, Stephen C., and Jacob C. Koella, eds. 2008. <em>Evolution in Health and Disease<\/em>. 2nd edition. United Kingdom: Oxford University Press.<\/p>\n<p>Zuk, Marlene. 2013. <em>Paleofantasy: What Evolution Really Tells Us about Sex, Diet, and <\/em><em>How We Live<\/em>. New York: W. W. Norton &amp; Company.<\/p>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Abid, Zaynah, Amanda J. Cross, and Rashmi Sinha. 2014. \u201cMeat, Dairy, and Cancer.\u201d <em>The American Journal of Clinical Nutrition<\/em> 100 (S1): 386S\u2013393S.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Akil, Luma, and H. Anwar Ahmad. 2011. \u201cRelationships between Obesity and Cardiovascular Diseases in Four Southern States and Colorado.\u201d <em>Journal of Health Care for the Poor and Underserved<\/em> 22 (S4): 61\u201372.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Ali, Aus Tariq. 2014. \u201cReproductive Factors and the Risk of Endometrial Cancers.\u201d <em>International Journal of Gynecological Cancer<\/em> 24 (3): 384\u2013393.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">American Heart Association. 2018. \u201cHeart Disease and Stroke Statistics-2018 Update: A Report.\u201d <em>Circulation 137<\/em> (12). Accessed April 7, 2023. https:\/\/www.ahajournals.org\/doi\/10.1161\/cir.0000000000000558.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">American Psychiatric Association (APA). 2013. <em>Diagnostic and Statistical Manual of Mental Disorder<\/em>. 5th Edition: DSM-5. Washington, DC: APA.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Armelagos, George J. 1990. \u201cHealth and Disease in Prehistoric Populations in Transition.\u201d <em>Disease in Populations in Transition: Anthropological and Epidemiological Perspectives,<\/em> edited by George J. Armelagos and Alan C. Swedland, 127\u2013144. New York: Bergin &amp; Garvey.<\/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 and Medicine 61<\/em> (4): 755-765.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0.495002746582031pt;margin-right: 0pt;text-indent: 0pt\">Arnot, Megan, Eva Brandl, O. L. K. Campbell, Yuan Chen, Mark Dyble, Emily H. Emmott, et al. 2020. <em>Evolution, Medicine, and Public Health<\/em> 2020 (1): 264\u2013278. https:\/\/doi.org\/10.1093\/emph\/eoaa038.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Asher, Claire. 2017. \u201cIllegal Bushmeat Trade Threatens Human Health and Great Apes.\u201d <em>Mongabay<\/em>, April 6. Accessed April 4, 2023. https:\/\/news.mongabay.com\/2017\/04\/illegal-bushmeat-trade-threatens-human-health-and-great-apes\/.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Baker, W. A., G. A. Hitman, K. Hawrami, M .I. McCarthy, A. Riikonen, E. Tuomilehto-Wolf, A. Nissinen, et al. 1994. \u201cApolipoprotein D Gene Polymorphism: A New Genetic Marker for Type 2 Diabetic Subjects in Nauru and South India.\u201d <em>Diabetic Medicine<\/em> 11 (10): 947\u2013952.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Baltic, Milan Z., and Marija Boskovic. 2015. \u201cWhen Man Met Meat: Meat in Human Nutrition from Ancient Times Till Today.\u201d <em>Procedia Food Science 5<\/em>: 6- 9.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Blue Cross Blue Shield Association (BCBSA). 2017. \u201cDiabetes and the Commercially Insured U.S. Population.\u201d <em>The Health of America Report<\/em>, August 1. 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N.d. \u201cGlobal Health - Pakistan.\u201d Accessed April 4, 2023. https:\/\/www.cdc.gov\/globalhealth\/countries\/pakistan\/default.htm.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Centers for Disease Control and Prevention (CDC). 2014. \u201cBreastfeeding Report Card: United States\/2014.\u201d Atlanta, GA: Centers for Disease Control and Prevention.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Centers for Disease Control and Prevention (CDC). 2020. <em>National Diabetes Statistics Report: Estimates of Diabetes and Its Burden in the United States<\/em>. Accessed April 7, 2023 from <a class=\"rId83\" href=\"https:\/\/diabetesresearch.org\/wp-content\/uploads\/2022\/05\/national-diabetes-statistics-report-2020.pdf\">https:\/\/diabetesresearch.org\/wp-content\/uploads\/2022\/05\/national-diabetes-statistics-report-2020.pdf<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Centers for Disease Control and Prevention (CDC). 2022. \u201cCDC Releases Updated Maps of America\u2019s High Levels of Inactivity.\u201d January 20. https:\/\/www.cdc.gov\/media\/releases\/2022\/p0120-inactivity-map.html.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Cer\u00f3n, Ella. 2017. \u201cHere\u2019s Every Oreo Flavor Ever Created.\u201d <em>TeenVogue.com<\/em>, June 19. 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Accessed April 4, 2023. https:\/\/www.pewresearch.org\/fact-tank\/2018\/04\/05\/a-record-64-million-americans-live-in-multigenerational-households\/.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Cooling, Laura. 2015. \u201cBlood Groups in Infection and Host Susceptibility.\u201d <em>Clinical Microbiology Reviews<\/em> 28 (3): 801\u2013870.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Croston, Glenn. 2012. \u201cThe Thing We Fear More Than Death: Why Predators Are Responsible for Our Fear of Public Speaking.\u201d <em>Psychology Today<\/em> blog, November 29. 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Rosalie, and Michael Zimmerman. 2010. \u201cCancer: An Old Disease, A New Disease or Something In Between?\u201d <em>Nature Reviews: Cancer<\/em> 10 (10): 728\u2013733.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Davis, Mike. 2005. <em>The Monster at Our Door: The Global Threat of Avian Flu<\/em>. New York: Owl Books.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">de Silva, A. M., K. R. Walder, T. J. Aitman, T. Gotoda, A. P. Goldstone, A. M. Hodge, M. P. de Courten, P. Z. Zimmet, and G. R. 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Boyd, Melvin Konner, and Marjorie Shostak. 1988. \u201cStone Agers in the Fast Lane: Chronic Degenerative Diseases in Evolut<\/span>ionary Perspective.\u201d <em>American Journal of Medicine<\/em> 84 (4): 739\u2013749.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Eaton, S. Boyd, Malcolm C. Pike, Roger V. Short, Nancy C. Lee, James Trussell, Robert A. Hatcher, James W. 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Robinson, Jad Farha, Amanda Bradke, Sherita H. Golden, et al. 2022. \u201cRacial and Ethnic Discrepancy in Pulse Oximetry and Delayed Identification of Treatment Eligibility among Patients With COVID-19.\u201d <em>JAMA Internal Medicine<\/em> 182 (7): 730\u2013738.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Flinn, Mark V., and Barry G. England. 2003. \u201cChildhood Stress: Endocrine and Immune Responses to Psychosocial Events.\u201d In <em>Social and Cultural Lives of Immune Systems: Theory and Practice in Medical Anthropology and International Health<\/em>, edited by James M. Wilce Jr., 105\u2013146. London: Routledge.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Food Empowerment Project. N.d. \u201cFood Deserts.\u201d Accessed April 4, 2023. https:\/\/foodispower.org\/access-health\/food-deserts\/.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Gao, George. 2015. \u201cAmericans\u2019 Ideal Family Size Is Smaller Than It Used to Be.\u201d Pew Research Center, May 8. Accessed April 4, 2023. https:\/\/www.pewresearch.org\/fact-tank\/2015\/05\/08\/ideal-size-of-the-american-family\/.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Ghosh, A. K., S. Venkatraman, O. Soroka, E. Reshetnyak, M. Rajan, A. An, J. K. 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Stains. 2012. \u201cObesity and Cancer Risk: Evidence, Mechanisms, and Recommendations.\u201d Special issue, \u201cNutrition and Physical Activity in Aging, Obesity, and Cancer,\u201d <em>Annals of the New York Academy of Sciences<\/em> 1271 (1): 37\u201343.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Walker, Polly, Pamela Rhubart-Berg, Shawn McKenzie, Kristin Kelling, and Robert S. Lawrence. 2005. \u201cPublic Health Implications of Meat Production and Consumption.\u201d <em>Public Health Nutrition<\/em> 8 (4): 348\u2013356.<\/p>\n<p class=\"import-Normal\">Washington, Harriet A. 2006. <em>Medical Apartheid: The Dark History of Medical Experimentation on Black Americans from Colonial Times to the Present.<\/em> New York: Anchor Books.<\/p>\n<p class=\"import-Normal\">Williams, David R., and Selina A. Mohammed. 2013. \u201cRacism and Health I: Pathways and Scientific Evidence.\u201d <em>American Behavioral Scientist<\/em> 57 (8). https:\/\/doi.org\/<a class=\"rId87\" href=\"https:\/\/doi.org\/10.1177%2F0002764213487340\">10.1177\/0002764213487340. <\/a><\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Williams, David R., and Ronald Wyatt. 2015. \u201cRacial Bias in Health Care and Health: Challenges and Opportunities.\u201d <em>JAMA <\/em>314 (6): 555\u2013556. https:\/\/doi.org10.1001\/jama.2015.9260.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Wolfe, Nathan, Claire P. Dunavan, and Jared Diamond. 2012. \u201cOrigins Of Major Human Infectious Diseases.\u201d In <em>Institute of Medicine: Improving Food Safety through a One Health Approach: Workshop Summary<\/em>, A16. Washington, DC: National Academies Press. Accessed April 4, 2023. https:\/\/www.ncbi.nlm.nih.gov\/books\/NBK114494\/.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">World Health Organization (WHO). 2016. <em>Global Report on Diabetes<\/em>. Accessed April 4, 2023. https:\/\/apps.who.int\/iris\/bitstream\/handle\/10665\/204871\/9789241565257_eng.pdf.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">World Health Organization (WHO). 2017. \u201cObesity and Overweight.\u201d Fact Sheet. Last modified June 9, 2021; accessed April 4, 2023. https:\/\/www.who.int\/mediacentre\/factsheets\/fs311\/en\/.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">World Health Organization (WHO). 2018. \u201cCancer.\u201d Fact Sheet. Last modified February 3, 2022; accessed April 5, 2023. https:\/\/www.who.int\/news-room\/fact-sheets\/detail\/cancer.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Worobey, Michael, Joshua I. Levy, Lorena Malpica Serrano, Alexander Crits-Christoph, Jonathan E. Pekar, Stephan A. Goldstein, Angela L. Rassmussen, et al. July 26, 2022. \u201cThe Huanan Seafood Wholesale Market in Wuhan was the Early Epicenter of the COVID-19 Pandemic.\u201d <em>SCIENCE <\/em>26 (377): 951\u2013959. <a class=\"rId88\" href=\"https:\/\/doi.org\/10.1126\/science.abp8715\">https:\/\/doi.org\/10.1126\/science.abp8715<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Worthman, Carol M., and Jennifer Kuzara. 2005. \u201cLife History and the Early Origins of Health Differentials.\u201d <em>American Journal of Human Biology<\/em> 17 (1): 95\u2013112.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Wrangham, Richard. 2009. <em>Catching Fire: How Cooking Made Us Human<\/em>. New York: Basic Books.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Yehuda, Rachel, and Joseph LeDoux. 2007. \u201cResponse Variation Following Trauma: A Translational Neuroscience Approach to Understanding PTSD.\u201d <em>Neuron<\/em> 56 (1): 19\u201332.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Zuckerman, Molly K., and George J. Armelagos. 2014. \u201cThe Hygiene Hypothesis and the Second Epidemiologic Transition.\u201d In <em>Modern Environments and Human Health: Revisiting the Second Epidemiologic Transition<\/em>, edited by Molly K. Zuckerman, 301\u2013320. Hoboken, NJ: Wiley-Blackwell.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Zuckerman, Molly Kathleen, Kristin Nicole Harper, Ronald Barrett, and George John Armelagos. 2014. \u201cThe Evolution of Disease: Anthropological Perspectives on Epidemiologic Transitions.\u201d Special issue, \u201cEpidemiological Transitions: Beyond Omran\u2019s Theory,\u201d <em>Global Health Action<\/em> 7 (1): 23303. https:\/\/doi.org\/10.3402\/gha.v7.23303.<\/p>\n<\/div>\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_1683_1700\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1700\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\">Joylin Namie, Ph.D., Truckee Meadows Community College<\/p>\n<p class=\"import-Normal\"><em>This chapter is a revision from <\/em><em>\"<\/em><a class=\"rId6\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-11\/\" target=\"_blank\" rel=\"noopener\"><em>Chapter 16: Contemporary Topics: Human Biology and Health<\/em><\/a><em>\u201d by Joylin Namie. In <\/em><a class=\"rId7\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>Explorations: An Open Invitation to Biological Anthropology, <\/em><\/a><a class=\"rId8\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"><em>first edition<\/em><\/a><em>, <\/em><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\">Learning Objectives<\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li>Describe the major transitions in patterns of disease that have occurred throughout human evolution.<\/li>\n<li>Describe what is meant by a \u201cmismatch\u201d between our evolved biology and contemporary lifestyles and how this is reflected in modern disease patterns.<\/li>\n<li>Explain how the human stress response can positively and negatively have an impact on health.<\/li>\n<li>Explain what a \u201csyndemic\u201d is and why the COVID-19 pandemic represents one.<\/li>\n<li>Describe the ways institutionalized racism and bias in the medical field contributed to different rates of exposure, differential treatment, morbidity, and mortality from COVID-19 for different ethnic groups in the United States.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p>When was the last time you needed to do research for an upcoming paper? I bet you started by looking for information online. How did you go about your search? Which websites looked promising? Which ones did not entice you to click past the home page? Once you found one you thought might be useful, how much time did you spend searching for information? At what point did you decide to leave that site and move on? I would wager money that you never once thought your behaviour had anything to do with human evolution, but it does.<\/p>\n<p class=\"import-Normal\">Although we may not often stop to think about it, our evolutionary past is reflected in many aspects of modern life. The ways we \u201cforage\u201d for information on the internet mimics the ways we once foraged for food during our several-million-year history as hunter-gatherers (Chin Et al. 2015). Humans are visual hunters (Lieberman 2006). We practice optimal foraging strategy, meaning we make decisions based on energy return for investment (McElroy &amp; Townsend 2009). When we search for information online, we locate a \u201cpatch,\u201d in this case a website or research article, then quickly scan the contents to discern how much of it is useful to us. Like our hominin ancestors, we spend more time in \u201cpatches\u201d with abundant resources and abandon sites quickly once we have exhausted the available goods. As with internet searches, our evolutionary past is also reflected in the kinds of landscapes we find appealing, the foods that taste good to us, why we break a sweat at the gym, and why we have to go to the gym at all (Bogin 1991; Dutton 2009; Lieberman 2015). Many of the health problems facing humans in the 21st century also have their beginnings in the millions of years we roamed the earth as foragers.<\/p>\n<h2 class=\"import-Normal\">Preagricultural Humans<\/h2>\n<h3 class=\"import-Normal\"><strong>Diet<\/strong><\/h3>\n<p class=\"import-Normal\">Humans may be the species with the longest list of nutritional requirements (Bogin 1991). This is due to the fact that we evolved in environments where there was a high diversity of edible species but low densities of any given species. <em>Homo sapiens sapiens<\/em> require 45\u201350 essential nutrients for growth, maintenance, and repair of cells and tissues. These include protein, carbohydrates, fats, vitamins, minerals, and water. As a species, we are (or were) physically active with high metabolic demands. We are also <strong>omnivorous<\/strong> and evolved to choose foods that are dense in essential nutrients. One of the ways we identified high-calorie resources in our evolutionary past was through taste, and it is no accident that humans find sweet, salty, fatty foods appealing.<\/p>\n<p class=\"import-Normal\">The human predisposition toward sugar, salt, and fat is innate (Farb &amp; Armelagos 1980). Receptors for sweetness are found in every one of our mouth\u2019s 10,000 taste buds (Moss 2013). Preference for sweet makes sense in an ancestral environment where sweetness signaled high-value resources like ripe fruits. Likewise, \u201cthe long evolutionary path from sea-dwelling creatures to modern humans has given us salty body fluids, the exact salinity of which must be maintained\u201d (Farb &amp; Armelagos 1980), drawing us to salty-tasting things. Cravings for fat are also inborn, with some archaeological evidence suggesting that hominins collected animal bones for their fatty marrow, which contains two essential fatty acids necessary for brain development (Richards 2002), rather than for any meat remaining on the surface (Bogin 1991).<\/p>\n<p class=\"import-Normal\">Bioarchaeological studies indicate Paleolithic peoples ate a wider variety of foods than many people eat today (Armelagos Et al. 2005; Bogin 1991; Larsen 2014; Marciniak &amp; Perry 2017). Foragers took in more protein, less fat, much more fibre, and far less sodium than modern humans typically do (Eaton, Konner, &amp; Shostak 1988). Changes in tooth and intestinal morphology illustrate that animal products were an important part of human diets from the time of <em>Homo erectus<\/em> onward (Baltic &amp; Boskovic 2015; Richards 2002; Wrangham 2009). Once cooking became established, it opened up a wider variety of both plant and animal resources to humans. However, the protein, carbohydrates, and fats preagricultural peoples ate were much different from those we eat today. Wild game lacked the antibiotics, growth hormones, and high levels of cholesterol and saturated fat associated with industrialized meat production today (Walker Et al. 2005). Wild game was also protein dense, providing only 50% of energy as fat (Lucock Et al. 2014). The ways meat is prepared and eaten today also have implications for disease.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Meats cooked well done over high heat and\/or over an open flame, including hamburgers and barbecued meats, are highly carcinogenic due to compounds formed during the cooking process (Trafialek &amp; Kolanowski 2014). Processed meats that have been preserved by smoking, curing, salting, or by adding chemical preservatives such as sodium nitrite (e.g., ham, bacon, pastrami, salami, and beef jerky) have been linked to cancers of the colon, lung, and prostate (Abid, Cross, &amp; Sinha 2014; Figure 17.1). Nitrites\/nitrates have additionally been linked to cancers of the ovaries, stomach, esophagus, bladder, pancreas, and thyroid (Abid, Cross &amp; Sinha 2014). In addition, studies analyzing the diets of 103,000 Americans for up to 16 years indicate that those who ate grilled, broiled, or roasted meats more than 15 times per month were 17% more likely to develop high blood pressure than those who ate meat fewer than four times per month, and participants who preferred their meats well done were 15% more likely to suffer from <strong>hypertension<\/strong> (high blood pressure) than those who ate their meats rare (Liu 2018). A previous study of the same cohort indicated \u201cindependent of consumption amount, open-flame and\/or high-temperature cooking for both red meat and chicken is associated with an increased risk of type 2 diabetes among adults who consume animal flesh regularly\u201d (Liu Et al. 2018). Although meat has been argued to be crucial to cognitive and physical development among hominins (Wrangham 2009), there has been an evolutionary trade-off between the ability to preserve protein through cooking and the health risks of cooked meat and chemical preservatives.<\/p>\n<figure style=\"width: 343px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/06\/image1-6.png\" alt=\"Consecutive circles outline categories of cancer risk with images of processed meats and red meat.\" width=\"343\" height=\"424\" \/><figcaption class=\"wp-caption-text\">Figure 17.1: Positive associations have been observed between meat consumption and some types of cancer. The International Agency for Research on Cancer (2018) categorized four groupings of cancer risk. The first group is labeled \"causes cancer\", and the second group \"probably causes cancer\". Group 1 includes processed meats such as bacon, salami and hot dogs. Group 2A includes red meat such as beef, pork and lamb. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-11\/\">Carcinogenic Meats (Figure 16.1)<\/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>. [Includes <a href=\"https:\/\/pngimg.com\/download\/10217\">Hot dog PNG image<\/a> by unknown, <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0<\/a>; <a href=\"https:\/\/www.publicdomainpictures.net\/en\/view-image.php?image=109418&amp;picture=rasher-of-bacon\">Rasher of Bacon<\/a> by unknown, <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/\">public domain (CC0)<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Salami_aka.jpg\">Salami aka<\/a> by Andr\u00e9 Karwath <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Aka\">Aka<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.5\/legalcode\">CC BY-SA 2.5<\/a>; <a href=\"https:\/\/pngimg.com\/download\/2127\">Cow PNG image<\/a> by unknown, <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0<\/a>; <a href=\"https:\/\/pngimg.com\/download\/2184\">sheep PNG image<\/a> by unknown, <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0<\/a>; <a href=\"https:\/\/www.publicdomainpictures.net\/en\/view-image.php?image=55516&amp;picture=pig-on-white-background\">Pig on white background<\/a> by unknown, <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/\">public domain (CC0)<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Although carbohydrates represent half of the diet on average for both ancient foragers and modern humans, the types of carbohydrates consumed are very different. Ancient foragers ate fresh fruits, vegetables, grasses, legumes, and tubers, rather than the processed carbohydrates common in industrialized economies (Moss 2013). Their diets also lacked the refined white sugar and corn syrup found in many modern foods that contribute to the development of diabetes (Pontzer Et al. 2012).<\/p>\n<h3 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Physical Activity Patterns<\/strong><\/h3>\n<p class=\"import-Normal\">How do we know how active our ancestors were? Hominin morphology and physiology provide us with clues. Adaptations to heat discussed in Chapter 10 evolved in response to the need for physical exertion in the heat of the day in equatorial Africa. Human adaptations for preventing hyperthermia (overheating) suggest an evolutionary history of regular, strenuous physical activity. Research with modern foraging populations also offers clues to ancient activity patterns. Although subject to sampling biases and marginal environments (Marlowe 2005), modern foragers provide the only direct observations of human behaviour in the absence of agriculture (Lee 2013). From such studies, we know foragers cover greater distances in single-day foraging bouts than other living primates, and these treks require high levels of cardiovascular endurance (Raichlen &amp; Alexander 2014). Recent research with the Hadza in Tanzania indicates they walk up to 11 kilometres (6.8 miles) daily while hunting and gathering (Pontzer Et al. 2012), engaging in moderate-to-vigorous physical activity for over two hours each day\u2014meeting the U.S. government\u2019s weekly requirements for physical activity in just two days (Raichlen Et al. 2016; Figure 17.2). The fact that humans were physically active in our evolutionary past is also supported by the fact that regular physical exercise has been shown to be protective against a variety of health conditions found in modern humans, including <strong>cardiovascular disease<\/strong> (Raichlen &amp; Alexander 2014) and Alzheimer\u2019s dementia (Mandsager, Harb, &amp; Cremer 2018), even in the presence of brain pathologies indicative of cognitive decline (Buchman Et al. 2019).<\/p>\n<p>&nbsp;<\/p>\n<figure style=\"width: 624px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image10-7.png\" alt=\"Two hunters with dogs and bows walk in a savannah. \" width=\"624\" height=\"417\" \/><figcaption class=\"wp-caption-text\">Figure 17.2: Hadza foragers hunting on foot. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Hadazbe_returning_from_hunt.jpg\">Hadazbe returning from hunt<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/7177420@N03\">Andreas Lederer<\/a> has been modified (cropped) and is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\">CC BY 2.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Infectious Disease<\/strong><\/h3>\n<p class=\"import-Normal\">Population size and density remained low throughout the Paleolithic, by some estimates only 0.4 inhabitants per square kilometre (McClellan &amp; Dorn 2006). This limited <strong>morbidity<\/strong> and <strong>mortality <\/strong>from infectious diseases, which sometimes require large populations to sustain epidemics. Our earliest ancestors had primarily two types of infections with which to contend (Armelagos 1990). The first were organisms that adapted to our prehominin ancestors and have been problems ever since. Examples include head lice, pinworms, and yaws. A second set of diseases were <strong>zoonoses<\/strong>, diseases that originate in animals and mutate into a form infectious to humans. One example is the Human Immunodeficiency Virus (HIV) that originated in nonhuman primates and was likely passed to humans through the butchering of hunted primates for food (Sharp &amp; Hahn 2011). Zoonoses that could have infected ancient hunter-gatherers include tetanus and <strong>vector-borne diseases<\/strong> transmitted by flies, mosquitoes, fleas, midges, ticks, and the like. Many of these diseases are slow acting, chronic, or latent, meaning they can last for weeks, months, or even decades, causing low levels of sickness and allowing victims to infect others over long periods of time. Survival or cure does not result in lasting immunity, with survivors returning to the pool of potential victims. Such diseases often survive in animal reservoirs, reinfecting humans again and again (Wolfe Et al. 2012). A study of bloodsucking insects preserved in samples of amber dating from 15 to 100 million years ago indicates that they carried microorganisms that today cause diseases such as river blindness, typhus, Lyme disease, and malaria (Poinar 2018). Such diseases may have been infecting humans throughout our history and may have had significant impacts on small foraging communities because they more often infected adults, who provided the food supply (Armelagos Et al. 2005).<\/p>\n<h3 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Health<\/strong><\/h3>\n<p class=\"import-Normal\">Given their diets, levels of physical activity, and low population densities, nomadic preagricultural humans were likely in better health than many modern populations. This assertion is supported by comparative research conducted with modern foraging and industrialized populations. Measures of health taken from 20th-century foraging populations demonstrate excellent aerobic capacity, as measured by oxygen uptake during exertion, and low body-fat percentages, with <strong>triceps skinfold measurements<\/strong> half those of white Canadians and Americans. Serum cholesterol levels were also low, and markers for diabetes, hypertension, and cardiovascular disease were missing among them (Eaton, Konner, &amp; Shostak 1988; Raichlen Et al<em>. <\/em>2016).<\/p>\n<h2 class=\"import-Normal\">Health Consequences of the Transition to Agriculture and Animal Domestication<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The shift from foraging to food production occurred relatively recently in our evolutionary history (Larsen 2014), and there are indications our biology has not yet caught up (Pritchard 2010). Beginning around 12,000 BCE in several parts of the globe, humans began to move toward a diet based on domesticated plants and animals (Armelagos Et al. 2005). This involved manipulating the natural landscape to facilitate intensive food production, including the clearing of forest and construction of wells, irrigation canals, and ditches, exposing humans to water-borne illnesses and parasites and attracting mosquitos and other vectors of disease to human settlements. The heavy, repetitive physical labour of early agricultural production resulted in negative impacts on articular joints, including <strong>osteoarthritis<\/strong> (Larsen 2014). At the same time, nutritional diversity became restricted, focused on major cereal crops that continue to dominate agricultural production today, including corn, wheat, and rice (Jain 2012). This represented a major shift in diet from a wide variety of plant and animal foods to dependence on starchy carbohydrates, leading to increases in dental caries (cavities), reductions in stature and growth rates, and nutritional deficiencies (Larsen 2014). Domesticated animals added new foods to the human diet, including meat that was higher in fat and cholesterol than wild game as well as dairy products (Lucock Et al. 2014). Agriculture provided the means to produce a storable surplus for the first time in human history, leading to the beginnings of economic inequality (see Chapter 13). Social hierarchies led to the unequal distribution of resources, concentrating infectious disease among the poor and malnourished (Zuckerman Et al. 2014), a situation that continues to plague humanity today (Marmot 2005).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\"><strong>Sedentism<\/strong> and a rise in population density accompanied the move to agriculture, increasing the risk of infectious disease. Agriculture often provided enough calories, if not enough nutrition, to increase fertility. Although diets were worse and people unhealthier, populations continued to grow, even in the midst of high levels of child and maternal mortality and short life expectancies (Omran 2005). Hygiene became an issue as large settlements increased the problem of removing human waste and providing uncontaminated water (Armelagos Et al. 2005). Domesticated animals provided reservoirs of zoonotic pathogens, which affected farmers more than foragers, as farmers were in closer proximity to their animals on a daily basis (Marciniak &amp; Perry 2017). Many of these diseases became major killers of humankind, including influenza, tuberculosis, malaria, plague, syphilis, and smallpox, functioning as selective pressures in and of themselves (Cooling 2015). As these diseases encountered large human populations, they caused major epidemics that traveled along newly established routes for trade, warfare, and colonization.<\/p>\n<h2 class=\"import-Normal\">Epidemiological Transitions<\/h2>\n<p class=\"import-Normal\">Changes in diet and physical-activity patterns, population densities, and exposure to zoonoses associated with agriculture resulted in an epidemiological transition, a shift in the causes of morbidity (sickness) and mortality (death) among humankind (Omran 2005). The first epidemiological transition from foraging to food production resulted in increases in dental caries (see Chapter 13), nutritional deficiencies, infectious disease, and skeletal conditions like osteoarthritis, as well as decreases in growth and height (Larsen 2014). A second epidemiological transition occurred following the Industrial Revolution in Western Europe and the United States when improved standards of living, hygiene, and nutrition minimized the effects of infectious disease, after which people began to experience higher rates of <strong>noncommunicable diseases<\/strong>, such as <strong>cancer<\/strong>, heart disease, and diabetes due to the changes in lifestyle, diet, and activity levels that are the subject of this chapter (Omran 2005). With the addition of immunizations and other public health initiatives, modified forms of this transition remain ongoing in many low- and middle-income countries (Zuckerman Et al. 2014), with several now facing a <strong>\u201cdouble burden\u201d <\/strong>of disease, with poor, often rural, populations struggling with infectious diseases due to malnutrition, lack of sanitation, and access to health care, while more affluent citizens are victims of chronic illnesses. A third epidemiological transition is now underway as infectious diseases, including new, re-emergent, and multidrug-resistant diseases, have once again become major health concerns (Harper &amp; Armelagos 2010; Zuckerman Et al. 2014). These include COVID-19, Ebola, HIV\/AIDS, tuberculosis, malaria, dengue, Lyme disease, and West Nile virus\u2014all zoonoses that initially spread to humans through contact with animals.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Patterns of morbidity and mortality continue to shift across the globe. As with the first epidemiological transition resulting from the adoption of large-scale agriculture, such shifts can be the direct, if unintended, result of human interactions with the environment. For example, there has been a rise in chronic inflammatory diseases (CIDs) in developed countries (Versini Et al<em>.<\/em> 2015). This includes increased rates of allergic conditions like asthma and autoimmune diseases like rheumatoid arthritis, multiple sclerosis, Crohn\u2019s disease, and inflammatory bowel disease. This has coincided with the decrease in infectious disease associated with the second epidemiological transition, and the two are related. The \u201chygiene hypothesis\u201d postulates the rise in CIDs is a result of limited exposure to nonlethal environmental pathogens in utero and early childhood (Zuckerman &amp; Armelagos 2014). Modern human societies have become so sanitized that we are no longer exposed to microorganisms that stimulate the development of a healthy immune system (Versini Et al. 2015). \u201cIn effect, the lifestyle changes\u2014sanitary improvements, pasteurization, use of antibiotics, and improved hygiene\u2014that contributed to the second transition may have produced a substantial trade-off, with developed nations exchanging a high burden of infectious disease for a higher burden of CIDs\u201d (Zuckerman Et al. 2014).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The third epidemiological transition, the re-emergence of infectious disease, reflects the continuing relationship between humans, animals, and pathogens. Over 60% of <strong>e<\/strong><strong>merging infectious diseases (EIDs)<\/strong> since 1940 have been of zoonotic origin, with over 70% stemming from human contact with wildlife (Jones Et al. 2008), including COVID-19. The crossover of COVID to humans is believed to have involved transmission from bats to an intermediate species then to humans, with infected humans then passing it to other humans in a wet market in Wuhan, China in late 2019 (Worobey Et al. 2022). Two COVID variants, representing two distinct crossover events from animals to humans, were circulating in the market by February 2020. Similarly, the global bushmeat trade currently devastating Africa\u2019s wildlife is a continuing source of Ebola infection (Asher 2017), as well as the original source of HIV and viruses related to leukaemia and lymphoma among humans (Zuckerman Et al. 2014). New strains of avian (bird) flu, some with mortality rates as high as 60% among humans (WHO n.d.), are transmitted to humans through poultry production and contact with wild birds (Davis 2005). Lastly, the use of antibiotics in commercial meat production is directly related to the rise of drug-resistant strains of previously controlled infectious diseases. An estimated 80% of antibiotics in the U.S. are used to promote growth and prevent infection in livestock, and drug-resistant bacteria from these animals are transmitted to humans through meat consumption (Ventola 2015).<\/p>\n<figure style=\"width: 299px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image13-7.png\" alt=\"Aerial photograph of a flooded city.\" width=\"299\" height=\"399\" \/><figcaption class=\"wp-caption-text\">Figure 17.3: Flooding in Sindh, Pakistan, in 2022. Credit: <a href=\"https:\/\/www.flickr.com\/photos\/193804179@N08\/52331043544\/\">Flood in Pakistan<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/193804179@N08\/\">Ali Hyder Junejo<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\">CC BY 2.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">A fourth epidemiological transition is currently underway in which some parts of the globe are suffering from a <strong>\u201ctriple burden\u201d<\/strong> of infectious and chronic diseases combined with injuries and diseases related to intensifying globalization, urbanization, deforestation, and climate change (Karn &amp; Sharma 2021). Massive flooding in Pakistan in 2022 (Figure 17.3) will serve to illustrate the concept. Following a severe heat wave in June 2022, Pakistan experienced extremely heavy seasonal monsoon rains, in some provinces 700% above normal. Combined with water flow from melting glaciers, this caused the worst flooding in the country\u2019s history, putting one third of the nation under water (Sheerazi 2022). The heat wave, glacial melt, and extreme rainfall were all attributable to global climate change, inflicting destruction and disease on Pakistan, which produces less than 1% of total global carbon emissions (Government of Pakistan 2021).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">As a direct result of the flooding, infrastructure, including roads, homes, and bridges, was destroyed, and 1,700 people died, nearly 13,000 were injured, and over 33 million were displaced. In addition to their initial injuries and trauma, displaced people lacked food, health care, safe water, and basic sanitation, leading to starvation and exposure to infectious diseases like malaria and dengue fever, as well as skin conditions like scabies, caused by mites. Pakistan also has a poverty rate of 30\u201340%, contributing to already-high rates of HIV, tuberculosis, and hepatitis. At the same time, the leading causes of death are heart disease, cancer, lower respiratory diseases, and stroke (CDC n.d), all chronic conditions.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">These examples illustrate continuing interaction between humans, our evolved biology, and the physical and cultural environments in which we live. The remainder of this chapter will focus on selected diseases and the social, cultural, and environmental factors that contribute to their <strong>prevalence<\/strong> in modern, industrialized economies. We begin with the health condition that affects all of the others\u2014<strong>obesity<\/strong>.<\/p>\n<h2 class=\"import-Normal\">Obesity<\/h2>\n<p class=\"import-Normal\">According to the World Health Organization (2017), 1.9 billion of the world\u2019s people are overweight and 650 million of these are obese. In the United States, 70% of Americans are overweight, and 40% of these meet the criteria for obesity. For the first time in human history, most of the world's population lives in countries where overweight and obesity kill more people than hunger (Figure 17.4). Improvements in public health and food production have allowed a greater number of people to live past childhood and to have enough to eat. This does not include everyone. Many people still struggle with poverty, hunger, and disease, even in the wealthiest of nations, including the United States. On a global scale, however, many people not only have enough food to survive but also to gain weight\u2014enough extra weight to cause health problems.<\/p>\n<p><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image4-5.png\" alt=\"&quot;Bar\" \/><\/p>\n<p class=\"import-Normal\">Although studies show differences in daily energy expenditure between foraging and farming populations compared with industrialized peoples, the major contributor to obesity in Western populations is energy intake (Pontzer Et al. 2012). Many people not only eat too much but too much of the wrong things. Biological anthropologist Leslie Lieberman (2006) argues that contemporary humans continue to rely on cues from foraging strategies of our evolutionary past that are now counterproductive in the <strong>obesogenic<\/strong> environments in which we now live.<\/p>\n<p class=\"import-Normal\">Examine your own eating habits in the context of how humans once hunted and gathered. Humans once relied on visual cues to find food, often traveled long distances to obtain it, then transported it back to our home base. There they may have had to process it by hand to render it edible. Think of how much less energy it takes to find food now. If we have the financial resources, we can acquire big energy payoffs by simply sitting at home and using an app on our mobile phone to place an order for delivery. And, voila! High-calorie (if not highly nutritious) food arrives at our door. Should we venture out for food, search time is reduced by signage and advertising directing us toward high-density \u201cpatches\u201d where food is available 24 hours a day. These include vending machines, gas stations, and fast-food outlets. Travel time is minimal and little human energy is used in the process (Lieberman 2006).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Foods are also prepackaged and prepared in ways that allow us to eat large quantities quickly. Think French fries or chicken nuggets, which we can easily eat with our hands while doing other things, like driving or watching television, rendering eating mindless and allowing us to take in food faster than our <strong>endocrine systems<\/strong> can tell us we are getting full. Modern \u201cpatches\u201d offer low-fibre, calorie-dense resources, which allow us to eat larger quantities, a problem already encouraged by larger portion sizes (Lieberman 2006). Processed foods are also engineered to appeal to human preferences for sweet tastes and fatty, creamy textures (Moss 2013). Remember from earlier chapters that natural selection favoured depth perception, colour vision, grasping hands, and coordinated eye-hand movements as general primate traits. Advertising and packaging now use our colour vision against us, attracting us to products that have little nutritional value but that play to our evolutionary predisposition toward variety. Remember those 50 different nutrients we require? That variety is now presented to us in the form of 55 different flavours of Oreo cookies (Cer\u00f3n 2017), which we take out of the package and dip in milk using our hand-eye coordination and depth perception.<\/p>\n<p class=\"import-Normal\">Even if we are ostensibly eating the same things our ancestors did, these foods are nothing alike. Take potatoes, for example. One medium-sized, plain, baked potato is a healthy food, especially if we eat the skin too. It contains 110 calories, 0 grams of fat, 26 grams of carbohydrates, and 3 grams of protein, plus 30% of the U.S. Recommended Daily Allowance (RDA) of vitamin C, 10% of vitamin B6, 15% of potassium, and no sodium (<a href=\"https:\/\/potatogoodness.com\/\" target=\"_blank\" rel=\"noopener\">Potato USA<\/a>). In contrast, a medium order of McDonald\u2019s fries, which takes the potato and adds salt and fat, contains 340 calories, 16 grams of fat, 44 grams of carbohydrates, 4 grams of protein, and 230 mg of sodium (<a href=\"https:\/\/www.mcdonalds.com\" target=\"_blank\" rel=\"noopener\">McDonalds<\/a>). Potato chips take food processing to a whole new level, removing even more nutrition and adding a host of additional ingredients, including oils, preservatives, and artificial flavourings and colours (Moss 2013). Take Ruffles Loaded Bacon and Cheddar Potato Skins Potato Chips as an example (St. Pierre 2018). The number of ingredients increases from one to 11 to 35 as we move from the potato to the potato chip, moving further from nature with each step (Figure 17.5). It should be noted that the nutritional information for the potato chips is based on a serving size of 11 chips, an amount likely smaller than many people eat. Many sweet, fatty, salty foods like fries and chips are cheap, which is why many people choose to eat them (Moss 2013). The price of a medium-sized order of McDonald\u2019s fries as of this writing is US$1.79, and the potato chips are $2.98 for an 8.5-ounce bag. A single potato prewrapped for microwaving is available in many supermarkets for US$1.99 but requires access to a microwave and eating utensils, making it less convenient.<\/p>\n<\/div>\n<div style=\"text-align: left\">\n<table class=\"grid\" style=\"border-collapse: collapse;width: 100%;height: 197px\" border=\"0\">\n<caption>Figure 17.5: The potato in three modern forms. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-11\/\">The potato in three modern forms (Figure 16.4)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Joylin Namie and 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>. [Includes <a href=\"https:\/\/www.publicdomainpictures.net\/en\/view-image.php?image=137873&amp;picture=potato\">Potato<\/a> by Charles Rondeau, <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/\">public domain (CC0)<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:McDonalds-French-Fries-Plate.jpg\">McDonalds-French-Fries-Plate<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Evan-Amos\">Evan-Amos<\/a>, <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/\">public domain (CC0)<\/a>; <a href=\"https:\/\/pdpics.com\/photo\/2316-potato-chips-bowl\/\">Potato chips bowl<\/a> by pdpics.com, <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/\">public domain (CC0)<\/a>.]<\/caption>\n<thead>\n<tr style=\"height: 47px\">\n<td style=\"width: 20.0464%;height: 47px\"><\/td>\n<td style=\"width: 25.4644%;height: 47px\">\n<p style=\"text-align: center\"><strong>Baked Potato<\/strong><br \/>\n[based, skin on, plain]<\/p>\n<\/td>\n<td style=\"width: 27.4768%;height: 47px;text-align: center\"><strong>French Fries<\/strong><br \/>\n[Medium order]<\/td>\n<td style=\"width: 27.0124%;height: 47px\"><strong>Potato Chips<\/strong><br \/>\n[1 oz. serving of 11 chips]<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr style=\"height: 15px\">\n<td style=\"width: 20.0464%;height: 15px\">Calories<\/td>\n<td style=\"width: 25.4644%;height: 15px;text-align: center\">110<\/td>\n<td style=\"width: 27.4768%;height: 15px;text-align: center\">340<\/td>\n<td style=\"width: 27.0124%;height: 15px\"><\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"width: 20.0464%;height: 15px\">Calories from fat<\/td>\n<td style=\"width: 25.4644%;height: 15px;text-align: center\">0<\/td>\n<td style=\"width: 27.4768%;height: 15px;text-align: center\">144<\/td>\n<td style=\"width: 27.0124%;height: 15px\"><\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"width: 20.0464%;height: 15px\">Fat<\/td>\n<td style=\"width: 25.4644%;height: 15px;text-align: center\">0g<\/td>\n<td style=\"width: 27.4768%;height: 15px;text-align: center\">16g<\/td>\n<td style=\"width: 27.0124%;height: 15px\"><\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"width: 20.0464%;height: 15px\">Carbohydrates<\/td>\n<td style=\"width: 25.4644%;height: 15px;text-align: center\">26g<\/td>\n<td style=\"width: 27.4768%;height: 15px;text-align: center\">44g<\/td>\n<td style=\"width: 27.0124%;height: 15px\"><\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"width: 20.0464%;height: 15px\">Protein<\/td>\n<td style=\"width: 25.4644%;height: 15px;text-align: center\">3g<\/td>\n<td style=\"width: 27.4768%;height: 15px;text-align: center\">4g<\/td>\n<td style=\"width: 27.0124%;height: 15px\"><\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"width: 20.0464%;height: 15px\">Sodium<\/td>\n<td style=\"width: 25.4644%;height: 15px;text-align: center\">0g<\/td>\n<td style=\"width: 27.4768%;height: 15px;text-align: center\">230mg<\/td>\n<td style=\"width: 27.0124%;height: 15px\"><\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"width: 20.0464%;height: 15px\">Dietary fibre<\/td>\n<td style=\"width: 25.4644%;height: 15px;text-align: center\">2g<\/td>\n<td style=\"width: 27.4768%;height: 15px;text-align: center\">4g<\/td>\n<td style=\"width: 27.0124%;height: 15px\"><\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"width: 20.0464%;height: 15px\">Sugars<\/td>\n<td style=\"width: 25.4644%;height: 15px;text-align: center\">1g<\/td>\n<td style=\"width: 27.4768%;height: 15px;text-align: center\">0g<\/td>\n<td style=\"width: 27.0124%;height: 15px\"><\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"width: 20.0464%;height: 15px\">Cholesterol<\/td>\n<td style=\"width: 25.4644%;height: 15px;text-align: center\">0g<\/td>\n<td style=\"width: 27.4768%;height: 15px;text-align: center\">0g<\/td>\n<td style=\"width: 27.0124%;height: 15px\"><\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"width: 20.0464%;height: 15px\">Ingredients<\/td>\n<td style=\"width: 25.4644%;height: 15px;text-align: center\">Potato<\/td>\n<td style=\"width: 27.4768%;height: 15px;text-align: center\">Potatoes, vegetable oil (canola oil), soybean oil, hydrogenated soybean oil, natural beef flavour (wheat and milk derivatives), citric acid (preservative), dextrose, sodium acid pyrophosphate (main colour), salt.<\/td>\n<td style=\"width: 27.0124%;height: 15px\">Potatoes, vegetable oil (sunflower, corn, and\/or canola oil), bacon and cheddar loaded potato skins seasoning (maltodextrin - made from corn) salt, cheddar cheese (milk, cheese cultures, salt enzymes), sour cream (cultured cream, skin milk), whey, dried onion, monosodium.<\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\">Not only have we transformed the food supply and our eating in ways that are detrimental to our health, but these changes have been accompanied by reductions in physical activity. <strong>Sedentarism<\/strong> is built into contemporary lifestyles. Think of how much time you spent sitting down today. Some of it may have been in class or at work, some may have been driving a car or perhaps binge-watching your favourite show, playing a video game, or checking in on social media. An inactive lifestyle is almost dictated by the digital age (Lucock Et al<em>.<\/em> 2014). Levels of physical activity in many countries are now so low that large portions of the population are completely sedentary, including one in five Americans (CDC 2022). For a species whose biology evolved in an environment where walking, lifting, and carrying were part of daily life, this is unhealthy and often leads to weight gain.<\/p>\n<h2 class=\"import-Normal\">Biology and Genetics of Weight<\/h2>\n<figure style=\"width: 344px\" class=\"wp-caption alignright\"><img class=\"\" src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image5-7.png\" alt=\"Four individuals of various ages walk alongside trees.\" width=\"344\" height=\"229\" \/><figcaption class=\"wp-caption-text\">Figure 17.6: Participants of a walk against diabetes and for general fitness around Nauru airport. Credit: <a href=\"https:\/\/www.flickr.com\/photos\/106853342@N04\/10709038046\">Participants of a walk against Diabetes and for general fitness around Nauru airport<\/a> by Lorrie Graham, <a href=\"https:\/\/www.flickr.com\/photos\/dfataustralianaid\/\">Department of Foreign Affairs and Trade<\/a>, is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\">CC BY 2.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Research indicates multiple genetic variants influence weight gain, and they are not spread evenly among human populations. Tuomo Rankinen and colleagues (2006) identified 127 genes associated with obesity, of which 22 contributed to weight gain. Claude Bouchard (2007) then identified five categories of obesity-promoting genotypes. These genotypes promote sedentarism, result in low metabolism, and lead to poor regulation of appetite, and a propensity to overeat. An example of the impact such genotypes can have in an environment of plenty is found among the population of the Micronesian island of Nauru. Historically, the island was geographically isolated and the food supply was unpredictable. These conditions favoured genotypes that promoted the ability to rapidly build up and store fat in times of food availability. In Nauruans, there are two genetic variants favouring weight gain and insulin resistance, and both are associated with obesity and type 2 diabetes. One variant is also associated with hypertension. One of these variants is also found in Pima Indians, who live in parts of Arizona and Mexico. In the Pima, this variant is associated with a high <strong>b<\/strong><strong>ody mass index (BMI) <\/strong> and type 2 diabetes, although it is not associated with the same outcomes in Japanese and British subjects (de Silva Et al<em>.<\/em> 1999). The other variant was analysed in Finnish and South Indian populations, neither of whom experienced the same outcome as Nauruans. This suggests these alleles may act as modifying genes for type 2 diabetes in some population groups (Baker Et al. 1994). Unfortunately, Nauruans are one of those groups. Eventually, they became wealthy through phosphate mining on the island, gaining access to a calorie-rich Western diet of imported foods and developing a sedentary lifestyle. This resulted in rates of type 2 diabetes as high as 30\u201340% in Nauruans over the age of 15, which became the leading cause of death (Lucock Et al. 2014), something Nauruans are taking seriously (Figure 17.6).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Factors other than biology influence which populations that carry a genetic predisposition to diabetes actually express it. The Onk Akimel O\u2019odham (Pima) of Arizona, for example, were seriously impacted by U.S. government policies that affected water rights, forcing the population away from subsistence farming to dependence on government handouts and convenience food. This resulted in a significant loss of physical activity, malnutrition, and obesity. The Onk Akimel O\u2019odham continue to experience hardship due to high rates of unemployment, poverty, and depression, sometimes made worse by alcoholism. In the absence of these pressures, the Onk Akimel O\u2019odham were diabetes free for centuries prior, even though they relied on agriculture for subsistence, suggesting genetics alone is not responsible for high rates of obesity and diabetes in current Onk Akimel O\u2019odham populations (Smith-Morris 2004).<\/p>\n<figure style=\"width: 372px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image3-4-1.jpg\" alt=\"A human body outline with multiple points, each listing related medical complications.\" width=\"372\" height=\"493\" \/><figcaption class=\"wp-caption-text\">Figure 17.7: Medical complications of obesity include stroke, sleep apnea, lung disease, liver disease, gallstones, cancer, heart disease, diabetes, pancreatitis, abnormal periods and infertility, arthritis, inflamed veins, and gout. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Medical_complications_of_obesity.png\">Medical complications of obesity<\/a> by the <a href=\"https:\/\/www.cdc.gov\/\">Centers for Disease Control and Prevention (CDC)<\/a> has been modified (colour changed and cancer list shortened) 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\">Obesity also has an epigenetic component. You learned about epigenetics in Chapter 4. With regard to obesity, epigenetics is counterintuitive in that mothers who do not take in enough calories during pregnancy often give birth to babies who grow up to be fat. What takes place is the fetus receives signals during pregnancy from its mother through the placenta and intrauterine environment about environmental conditions outside of the womb, in this case food insecurity. These signals encourage the turning on and off of genes related to metabolism. This alters the phenotype of the fetus so that if the child is born into an environment where food is plentiful, it will put on weight rapidly whenever possible, leading to obesity and related diseases later in life. If the child is a girl, her own eggs are formed in utero with the same genetic changes coded in, meaning she will pass along this same genetic predisposition to gain weight to her children. Hence, a biological propensity toward obesity can continue across generations (Worthman &amp; Kuzara 2005). Epigenetic changes to genes that promote weight gain are argued to be partly responsible for the rapid rise in obesity and diabetes in developing countries gaining access to Western diets (Stearns, Nesse, and Haig 2008).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Obesity and overweight put a strain on several biological systems of the body, including the <strong>circulatory<\/strong>, endocrine, and skeletal systems, contributing to hypertension, heart disease, <strong>stroke<\/strong>, diabetes, and osteoarthritis (Figure 17.7). Obesity also elevates the risk of cancers of the breast, endometrium, kidney, colon, esophagus, stomach, pancreas, and gallbladder (National Institutes of Health 2017; Vucenik &amp; Stains 2012). Diabetes\u2014one of the fastest-growing health conditions around the globe (WHO 2016) and one tightly connected to obesity and overweight\u2014is the focus of the following Special Topics box.<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Diabetes<\/h2>\n<p class=\"import-Normal\"><strong>Diabetes mellitus<\/strong> is an endocrine disorder characterized by excessively high blood glucose levels (Martini Et al<em>.<\/em> 2013). According to a report released by the World Health Organization, the number of people living with diabetes is growing in all regions of the world. Rates of diabetes have nearly doubled in the past three decades, largely due to increases in obesity and sugary diets (WHO 2016). One in 10 people around the world, 537 million people, now have diabetes, and three out of four live in low- and middle-income countries (IDF 2022). In the United States, 37 million people have diabetes (CDC 2020), where the disease is rising fastest among millennials (those ages 20\u201340) (BCBSA 2017), and one in every two adults with diabetes is undiagnosed (IDF 2022). Obesity and diabetes are linked: obesity causes a diet-related disease (diabetes) because of humans\u2019 evolved metabolic homeostasis mechanism, which is poorly suited to contemporary energy environments (Lucock Et al<em>.<\/em> 2014).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">To function properly, cells need a steady fuel supply. Blood sugar (glucose) is the fuel for most cells in the body, and the body produces the hormone <strong>insulin<\/strong> to help move glucose into cells that need it (Figure 17.8). Foods that most readily supply glucose to your bloodstream are carbohydrates, especially starchy foods like potatoes or sweet, sugary foods like candy and soda. The body can also convert other types of foods, including protein-rich foods (e.g., lean meats) and fatty foods (e.g.<em>,<\/em> vegetable oils and butter), into blood sugar in the liver via gluconeogenesis. Insulin\u2019s main job is to tell your cells when to take up glucose. The cell also has to listen to the signal and mobilize the glucose transporters. This not only allows your cells to get the energy they need, but it also keeps blood sugar from building up to dangerously high levels when you are at rest.<\/p>\n<figure style=\"width: 485px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image7-7.png\" alt=\"The cycle of how a body with diabetes processes nutrients from food.\" width=\"485\" height=\"648\" \/><figcaption class=\"wp-caption-text\">Figure 17.8: Carbohydrates are eaten and broken down into simple sugars (e.g., glucose). Glucose enters the bloodstream from the intestines, and the increase in glucose stimulates the pancreas to release insulin into the bloodstream. Insulin deposits glucose in the muscles and fat cells, where it is stored and used for energy. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-11\/\">Glucose metabolism (Figure 16.7)<\/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\">This system has limits. Like the rest of our biology, it evolved during several million years when sugar was hard to come by and carbohydrates took the form of fresh foods with a low <strong>glycemic index (GI)<\/strong>. Our ancestors were also active throughout the day, taking pressure off of the endocrine system. Now, sedentary lifestyles and processed-food diets cause many of us to take in more calories\u2014and especially more carbohydrates\u2014than our bodies can handle. There is only so much blood sugar your cells can absorb. Many modern populations are taxing those limits. After years of being asked by insulin to take in more glucose than they can use, cells eventually stop responding (McKee &amp; McKee 2015). This is called type 2 diabetes or insulin resistance, which accounts for 90\u201395% of diabetes cases in the United States (Figure 17.9). type 1 diabetes is believed to be caused by an autoimmune response in which your immune system is attacking and destroying the insulin-producing cells in your pancreas (Figure 17.9). type 1 diabetes is a genetic condition that often shows up early in life, while type 2 is more lifestyle-related and develops over time.<\/p>\n<figure style=\"width: 462px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image9-6.png\" alt=\"Type 1 cells and Type 2 cells and their response to insulin.\" width=\"462\" height=\"276\" \/><figcaption class=\"wp-caption-text\">Figure 17.9: Type 1 and type 2 diabetes. For Type 1 Diabetes, cells do not absorb glucose becuase there is no insulin. For Type 2 Diabetes, although there is insulin available, cells do not respond to it. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-11\/\">Type 1 and Type 2 Diabetes (Figure 16.8)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson has been modified (text) and 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\">Cardiovascular Disease<\/h2>\n<p class=\"import-Normal\">Cardiovascular disease (CVD)\u2014which includes coronary heart disease, hypertension (high blood pressure), and stroke\u2014is the leading cause of death globally, and heart disease remains the number one cause of death in the United States (American Heart Association 2018). Risk factors for cardiovascular disease include diet, obesity\/overweight, diabetes, smoking and alcohol consumption, and physical inactivity.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The connections between these factors and heart disease may not seem obvious and will be addressed here beginning with diet. Diets high in saturated fat and cholesterol can lead to atherosclerosis, a condition in which fat and cholesterol form plaque inside the arteries, eventually building up and hardening to the point that blood flow is blocked. Too much salt in the diet leads to fluid retention, which increases blood volume and thereby blood pressure, taxing the heart. Obesity\/overweight contribute to cardiovascular disease directly through increases in total blood volume, cardiac output, and cardiac workload. In other words, the heart has to work much harder if one is overweight (Akil &amp; Ahmad 2011). Obesity also relates to CVD indirectly through elevation of blood pressure (hypertension) and diabetes. High levels of blood glucose from diabetes can damage blood vessels and the nerves that control the heart and blood vessels. Alcohol consumption can raise blood pressure and triglyceride levels, a type of fat found in the blood. Alcohol also adds extra calories, which may cause weight gain, especially around the abdomen, which is directly associated with risk of a heart attack (Akil &amp; Ahmad 2011). Cigarette smoking also increases the risk of coronary heart disease. Nicotine increases blood pressure; in addition, cigarette smoke causes fatty buildup in the main artery in the neck and thickens blood, making it more likely to clot. It also decreases levels of HDL (\u201cgood\u201d) cholesterol (American Heart Association 2018). Even secondhand smoke can have an adverse effect if exposure occurs on a regular basis. Chronic psychological stress also elevates the risk of heart disease (Dimsdale 2008). The repeated release of stress hormones like adrenaline elevates blood pressure and may eventually damage artery walls. The human <strong>stress response<\/strong> and its connections to health and disease are discussed in more detail below.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">However, physical activity alters the likelihood of having heart disease, both directly and indirectly. Regular exercise of moderate to vigorous intensity strengthens the heart muscle and allows capillaries, tiny blood vessels in your body, to widen, improving blood flow. Regular exercise can also lower blood pressure and cholesterol levels and manage blood sugar levels, all of which reduce the risk of CVD.<\/p>\n<h2 class=\"import-Normal\">Cancer<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Cancer is the second-leading cause of death globally, causing one in every six deaths and killing nearly nine million people in 2015 (WHO 2018). Lifetime cancer risk in developed Western populations is now one in two, or 50% (Greaves 2015). Approximately one-third of deaths from cancer are due to behavioural and dietary factors, including high body mass index (BMI), low fruit and vegetable intake, lack of physical activity, and the use of tobacco and alcohol. Depending on the type of cancer and one\u2019s own genetic inheritance, these factors can increase cancer risk from 2- to 100-fold (Greaves 2015). Cancer is the result of interactions between a person's genes and three categories of external agents: physical carcinogens (e.g., ultraviolet radiation), chemical carcinogens (e.g., tobacco smoke, asbestos), and biological carcinogens, such as infections from certain viruses, bacteria, or parasites (WHO 2018). Obesity is also a risk factor for cancer, including of the breast, endometrium, kidney, colon, esophagus, stomach, pancreas, and gallbladder (National Institutes of Health 2017; Vucenik &amp; Stains 2012).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Cancer has been regarded as a relatively recent affliction for humans that became a problem after we were exposed to modern carcinogens and lived long enough to express the disease (David &amp; Zimmerman 2010). Given the long history that humans share with many oncogenic (cancer-causing) parasites and viruses (Ewald 2018), and the recent discovery of cancer in the metatarsal bone of a 1.8-million-year-old hominin (Odes Et al. 2016), this view is being challenged (See \u201cSpecial Topic: Life Choices and Reproductive Cancers in Women\u201d). The difficulties of identifying cancer in archaeological populations are many. Most cancer occurs in soft tissue, which rarely preserves, and fast-growing cancers would likely kill victims before leaving evidence in bone. It is also difficult to distinguish cancer from benign growths and inflammatory disease in ancient fossils, and there is often postmortem damage to fossil evidence from scavenging and erosion. However, using 3-D images, South African researchers recently diagnosed a type of cancer called osteosarcoma in a toe bone belonging to a human relative who died in Swartkrans Cave between 1.6 and 1.8 million years ago (Randolph-Quinney Et al. 2016). This study provides the earliest evidence of cancer in hominins.<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Life Choices and Reproductive Cancers in Women<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Behavioural or \u201clifestyle\u201d choices have an impact on cancer risk. Breast cancer is one example. It is the most common cancer in women worldwide, but <strong>incidence<\/strong> of new cases varies from 19.3 per 100,000 women in Eastern Africa to 89.7 per 100,000 women in Western Europe (WHO 2018). These differences are attributable to cultural changes among women in Western, industrialized countries that are a mismatch for our evolved reproductive biology. Age at <strong>menarche<\/strong>, the onset of menstrual periods, has dropped over the course of the last century from 16 to 12 years of age in the U.S. and Europe, with some girls getting their periods and developing breasts as young as eight years old (Greenspan &amp; Deardorff 2014, Figure 17.10). A World Health Organization study involving 34 countries in Europe and North America suggests the primary reason for the increase in earlier puberty is obesity, with differences in BMI accounting for 40% of individual- and country-level variance (Currie Et al<em>.<\/em> 2012). Early puberty in girls is associated with increased risk of breast cancer, ovarian cancer, diabetes, and high cholesterol in later life (Pierce &amp; Hardy 2012).<\/p>\n<figure style=\"width: 554px\" class=\"wp-caption aligncenter\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image12-3.jpg\" alt=\"A graph shows the decrease in age at menarche for five European nations and United States.\" width=\"554\" height=\"434\" \/><figcaption class=\"wp-caption-text\">Figure 17.10: Decreasing ages at time of first menstruation in selected countries. Credit: <a href=\"https:\/\/en.wikipedia.org\/wiki\/Menarche#\/media\/File:Acceleration1.jpg\">Acceleration1.jpg<\/a> by Yahadzija 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>&nbsp;<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">At the same time that age at puberty is dropping for girls in Western nations, age at birth of the first child is later, at 26 years old (Mathews and Hamilton 2016). Women are also having fewer children, two on average (Gao 2015), with 15% of women choosing to remain childless (Livingston 2015). Rates of breastfeeding have risen in recent decades but drop to only 27% of infants once babies reach 12 months of age (CDC 2014). In contrast, data from modern foraging populations (Eaton Et al. 1994) indicate that age at menarche is around 16 years old, age at birth of the first child is 19, breastfeeding on demand continues for three years for each child, and the number of children averages six. These differences relate to elevated risk for reproductive cancers, including breast cancer, among women in developed countries.<\/p>\n<p class=\"import-Normal\">Other than an established genetic risk (e.g., BRCA gene), the primary risk factor for breast cancer is exposure to estrogen. For women living in modern, industrialized economies, this exposure now often comes from women\u2019s own ovaries rather than from external environmental sources (Stearns, Nesse, and Haig 2008). Women in cultures without contraception are pregnant or breastfeeding for much of their reproductive lives, resulting in 100 or so menstrual cycles per lifetime. In contrast, Western women typically experience 400 or more (Strassmann 1997). This is partly due to early puberty. From menarche to the birth of a woman\u2019s first child can be 14 years or longer in Western populations, after which breastfeeding, if undertaken at all, lasts for a few weeks or months. Oral contraceptives or other hormonal methods to control reproduction induce monthly periods. Age at menopause (the cessation of menstrual cycles) is 50\u201355 years old across human populations. For Western women, this translates into forty years of menstrual cycling. Each month, the body prepares for a pregnancy that never occurs, experiencing cell divisions that put women at risk for cancers of the breast, endometrium, ovaries, and uterus (Strassmann 1999). Obesity adds to the risk, as adipose (fat) tissues are the primary source of estrogen biosynthesis. Thus, weight gain during the postmenopausal stage means higher exposure to estrogen and a greater risk of cancer (Ali 2014).<\/p>\n<p class=\"import-Normal\">Women cannot return to our evolutionary past, and there are significant social and economic reasons for delaying pregnancy and having fewer children. These include achieving educational and career goals, greater earning power, a reduction in the gender pay gap, more enduring marriages, and a decrease in the number of women needing public assistance (Sonfield Et al. 2013). There are also cultural means by which we might reduce the risk of reproductive cancers. These include reformulating hormonal contraceptives with enough estrogen to maintain bone density but reducing the number of menstrual periods over the reproductive lifespan (Stearns, Nesse, &amp; Haig 2008). Reducing fat intake may also lower estrogen levels. High-fat diets contribute to breast tumor development, while high fibre diets are beneficial in decreasing intestinal resorption of estrogenic hormones. Exercise also appears protective. Studies of former college athletes demonstrate risks of breast, uterine, and ovarian cancers later in life, two to five times lower than those of nonathletes (Eaton Et al<em>.<\/em> 1994).<\/p>\n<\/div>\n<h2 class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Stress<\/h2>\n<p class=\"import-Normal\">Have you ever been \u201cstressed out\u201d in class? Say you\u2019re in a large lecture hall with a hundred other people, or even in a small class where you don\u2019t know anyone. You\u2019re not sure about something the professor just said and you would really like to ask about it, so you start to raise your hand. Does your heart begin to pound and your mouth become dry? Do you get so nervous that you choose to ask a classmate after lecture instead? If so, you are not alone. Fear of speaking in public is one of the most common social phobias (APA 2013). It has been estimated that 75% of all people experience some degree of anxiety or nervousness when it comes to public speaking (Hamilton 2011), and surveys have shown that most people fear public speaking more than they fear death (Croston 2012).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">We have evolution to thank for this.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Humans, like other primates, are social animals. Being part of a group helped us to survive predation, get enough to eat, and successfully raise our young. When faced with standing up in front of a group, or even speaking up in class, we break into a sweat because we are afraid of rejection. Psychologist Glenn Croston (2012) writes, \u201cThe fear is so great because we are not merely afraid of being embarrassed or judged. We are afraid of being rejected from the social group, ostracized and left to defend ourselves all on our own. We fear ostracism still so much today it seems, fearing it more than death, because not so long ago getting kicked out of the group probably really was a death sentence.\u201d Hence, it is no surprise that public speaking triggers a stress response among much of humankind.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The human nervous system evolved in a context where quick responses to perceived threats presented an evolutionary advantage. The \u201cfight or flight\u201d response was honed during millions of years when threats more often took the form of an approaching lion than an approaching deadline. Our body\u2019s stress response, however, is triggered by a wide variety of stressors that produce the same general pattern of hormonal and physiological adjustments (Martini Et al. 2013). In today\u2019s world, the system is often stuck in the \u201con\u201d position due to the constant pressures of modern life, and this is a significant influence on health and disease.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">It is important to recognize that there are different types of stress and the time in life when adult coping mechanisms are formed is in childhood. In children, some stressors can be positive\u2014for example, stressors that are mild to moderate in magnitude, and accompanied by the support of a caring adult, which help children develop pathways by which stress is dealt with by the body throughout life. In a young child, a positive stress response might be going to the pediatrician to receive a vaccination and receiving encouragement and comfort from both parent and practitioner. A tolerable stress response is more serious, precipitated by something like a divorce or death of a relative. Again, buffered by positive support from surrounding adults, these types of stressors can be successfully managed by children. Toxic stress, however, \u201cresults from strong, frequent or prolonged activation of the stress response in the absence of the buffering protection of a supportive adult relationship\u201d (Shonkoff &amp; Garner 2012). Examples include child abuse or neglect, parental substance abuse, homelessness, and violence. In the absence of adequate psychological and physical support, the biological pathways of a child\u2019s physiological stress response are altered and lead to reduced abilities to cope with life\u2019s challenges as an adult.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The negative effects of sustained, elevated cortisol levels on health are well documented. These include higher levels of infectious disease and slowed growth in childhood (Flinn &amp; England 2003) and increased incidence of heart disease, obesity, and diabetes in adults (Worthman &amp; Kuzara 2005). Contrary to our evolutionary past, many causes of sustained stress in contemporary societies are psychosocial rather than physical threats. These can include an unhappy marriage or frustrations at work (Dimsdale 2008). Stressors can also be more subtle. For example, a review of research into the effects of stress on health indicated that experiencing racism was a significant stressor that was associated with alcohol consumption, psychological distress, overweight, abdominal obesity, and higher fasting-glucose levels among minority groups (Williams &amp; Mohammed 2013). Chronic, everyday racial discrimination is also associated with the hardening of coronary arteries, elevated blood pressure, giving birth to lower-birth-weight infants, cognitive impairment, poor sleep, and visceral fat, which is fat stored deep inside the belly, wrapped around the organs, including the liver and intestines. Visceral fat is a sign of <strong>m<\/strong><strong>etabolic syndrome<\/strong>, increasing the risk of stroke, heart disease, and type 2 diabetes. These effects have been shown to increase morbidity and mortality among members of affected groups.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Epigenetics can also be a factor in how a person is able to deal with stressful situations. Maternal experiences of stress during pregnancy have the potential to permanently alter the physiology of mothers\u2019 offspring, especially the hypothalamic-pituitary-adrenal (HPA) axis. The HPA axis regulates metabolism, blood pressure, and the immune response, and these alterations can predispose prenatally stressed individuals to suffer metabolic, cardiovascular, and mental disorders in adulthood (Palma-Gudiel Et al. 2015). These experiences carry across generations, with children of Holocaust survivors who experienced PTSD demonstrating similar changes in neurochemistry in the absence of a sustained, traumatic event, as did infant offspring of mothers who developed PTSD during pregnancy after witnessing the traumatic events of 9\/11 (Yehuda &amp; LeDoux 2007).<\/p>\n<h2 class=\"import-Normal\" style=\"text-indent: 0pt\">Syndemics and the Ecological Model<\/h2>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">It is important to recognize that disease risk is not spread evenly within or between populations. Diseases combine and interact to create a <strong>syndemic<\/strong>, where the coexistence of two or more conditions exacerbates the effects of one or all conditions. A syndemic (versus a pandemic, for example) takes into account social, political, economic, and environmental factors that increase risk for the clustering of two or more diseases (Singer Et al. 2017). One of the first syndemics identified involved substance abuse, violence, and AIDS. In inner cities in the U.S., the health crisis around HIV\/AIDS was related to tuberculosis, sexually transmitted infections, hepatitis, cirrhosis, infant mortality, drug abuse, suicide, and homicide. These were connected to poverty, homelessness, unemployment, poor nutrition, lack of social support, and social and ethnic inequality (Singer Et al. 2017). Together, these factors and others, like health policy and unequal access to health care, form an <strong>ecological model<\/strong> of health and disease, one that moves beyond biology and medical intervention (Sallis Et al. 2008).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">The COVID-19 pandemic represents a syndemic in which systemic racism in the healthcare system, differential access to diagnosis and treatment, income, employment, housing, family structure, pre existing conditions, and public health policies combined to result in higher rates of infection and death for African Americans, Native Americans, Asians, and Hispanic populations in the United States (Figure 17.11).<\/p>\n<table class=\"grid\" style=\"border-collapse: collapse;width: 100%\" border=\"0\">\n<caption>Figure 17.11: Risk for COVID-19 infection, hospitalization, and death by race\/ethnicity. Race and ethnicity are risk markers for other underlying conditions that affect health, including socioeconomic status, access to health care, and exposure to the virus related to occupation, e.g., frontline, essential, and critical infrastructure workers. Credit: <a href=\"https:\/\/www.cdc.gov\/coronavirus\/2019-ncov\/covid-data\/investigations-discovery\/hospitalization-death-by-race-ethnicity.html\">Risk for COVID-19 Infection, Hospitalization, and Death by Race\/Ethnicity<\/a> by the <a href=\"https:\/\/www.cdc.gov\/\">Centers for Disease Control and Prevention<\/a> is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/caption>\n<thead>\n<tr class=\"shaded\">\n<td style=\"width: 130.367px\">Rate ratios compared to White, Non-Hispanic persons<\/td>\n<td style=\"width: 130.367px\">American Indian or Alaska Native, Non-Hispanic persons<\/td>\n<td style=\"width: 130.367px\">Asian, Non-Hispanic persons<\/td>\n<td style=\"width: 130.383px\">Black or African American, Non-Hispanic persons<\/td>\n<td style=\"width: 130.35px\">Hispanic or Latino persons<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr>\n<td style=\"width: 130.367px\">Cases<\/td>\n<td style=\"width: 130.367px\">1.6x<\/td>\n<td style=\"width: 130.367px\">.8x<\/td>\n<td style=\"width: 130.383px\">1.1x<\/td>\n<td style=\"width: 130.35px\">1.5x<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 130.367px\">Hospitalization<\/td>\n<td style=\"width: 130.367px\">2.7x<\/td>\n<td style=\"width: 130.367px\">.8x<\/td>\n<td style=\"width: 130.383px\">2.3x<\/td>\n<td style=\"width: 130.35px\">2.0x<\/td>\n<\/tr>\n<tr>\n<td style=\"width: 130.367px\">Death<\/td>\n<td style=\"width: 130.367px\">2.1x<\/td>\n<td style=\"width: 130.367px\">.8x<\/td>\n<td style=\"width: 130.383px\">1.7x<\/td>\n<td style=\"width: 130.35px\">1.8x<\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<p class=\"import-Normal\">COVID-19 was the third leading cause of death in the U.S. in 2020 and 2021 (NIH 2022; Figure 17.12), but morbidity and mortality was not equally spread across the population. Working-class people and people of colour in the U.S. are more likely to live in poverty, in areas with high rates of crime and violence, and in close proximity to freeways and environmental threats like petrochemical plants and waste incinerators (Singer and Baer 2012). Many such neighbourhoods are also food \u201cdeserts\u201d without ready access to a healthy, affordable diet, made more challenging by residents not owning a car (Food Empowerment Project n.d.). Low-income people also often lack access to high-quality health care and delay or avoid preventive care and health screenings (Ross Et al. 2007). These factors contributed to higher rates of preexisting conditions, including obesity, diabetes, hypertension, asthma, heart disease, chronic obstructive pulmonary disease (COPD), and smoking behaviour, which then led to more complications and higher death rates from COVID (Ghosh Et al. 2021).<\/p>\n<p class=\"import-Normal\">Family structure also affected COVID exposure and severity. Many Americans live in multigenerational households, including 27% of Hispanics, 29% of Asians, 26% of African Americans, and 20% of Whites (Cohn &amp; Passel 2018). Not all multigenerational households are equal, however. Over twice as many African Americans as Whites are in multigenerational families in which at least one family member is unemployed, and over three times as many African Americans are in multigenerational families in which everyone is simultaneously unemployed (Park, Wiemers, and Seltzer 2019). Family members in multigenerational households were at a much higher risk of developing more severe forms of COVID due to decreased personal space and multiple exposures to the virus, as well as higher rates of diabetes, smoking, and residents living below the poverty line (Ghosh Et al. 2021). While aimed at reducing overall infection rates from COVID, public health measures such as mandatory lockdowns only exacerbated the situation in overcrowded and multigenerational housing, resulting in higher rates of infection and death in these communities.<\/p>\n<div style=\"margin: auto\">\n<table class=\" aligncenter\" style=\"width: 468pt;height: 195px\">\n<caption>Figure 17.12: Top five causes of death in the U.S. and worldwide since 2020. Credit: Top five causes of death in the U.S. and worldwide original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Joylin Namie is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. Based on data from Shiels Et al. 2022 and Traeger 2022.<\/caption>\n<thead>\n<tr class=\"shaded\" style=\"height: 30px\">\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 164.6px\">\n<p class=\"import-Normal\"><strong>United <\/strong><strong>States<\/strong><\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 429.733px\">\n<p class=\"import-Normal\"><strong>Worldwide<\/strong><\/p>\n<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"Table2-R\" style=\"height: 0\">\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 164.6px\">\n<p class=\"import-Normal\">1. Heart disease<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 429.733px\">\n<p class=\"import-Normal\">1. Heart disease<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 0\">\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 164.6px\">\n<p class=\"import-Normal\">2. Cancer<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 429.733px\">\n<p class=\"import-Normal\">2. Stroke<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 0\">\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 164.6px\">\n<p class=\"import-Normal\">3. COVID-19<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 429.733px\">\n<p class=\"import-Normal\">3. COVID-19<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 0\">\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 164.6px\">\n<p class=\"import-Normal\">4. Accidents<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 429.733px\">\n<p class=\"import-Normal\">4. Chronic Obstructive Pulmonary Disease<\/p>\n<\/td>\n<\/tr>\n<tr class=\"Table2-R\" style=\"height: 0\">\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 164.6px\">\n<p class=\"import-Normal\">5. Stroke<\/p>\n<\/td>\n<td class=\"Table2-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 429.733px\">\n<p class=\"import-Normal\">5. Lower respiratory infections<\/p>\n<\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"height: 15px;width: 165.467px\"><\/td>\n<td style=\"height: 15px;width: 430.6px\"><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n<p class=\"import-Normal\">There is a long history of systemic racism and discrimination in the medical system in the United States (Washington 2006). African Americans have been subjected to medical testing and experimentation without their consent or knowledge since the time of slavery. They continue to routinely receive care of poorer quality than whites (Williams &amp; Wyatt 2015), less pain medication during treatment and hospitalization (Green Et al. 2003), and differential treatment during pregnancy and childbirth (Washington 2006). Many Americans, including 50% of White medical students and residents in one recent study (Hoffman Et al. 2016), hold at least one false belief about African Americans, including \u201cBlack people\u2019s skin is thicker than white people\u2019s skin,\u201d \u201cBlacks have stronger immune systems than whites,\u201d and \u201cBlacks\u2019 nerve endings are less sensitive than whites\u2019.\u201d Such beliefs affect health care for African Americans in medical emergencies and for chronic conditions.<\/p>\n<p class=\"import-Normal\">During the COVID-19 pandemic, patients with darker skin in the United States were negatively affected by the very medical device most commonly used to assess oxygen levels in their blood. The pulse oximeter, a small device that clips onto the tip of your index finger and measures blood oxygen levels, experienced increased use in home, clinical, and hospital settings during the COVID-19 pandemic. Decisions regarding treatment and hospital admission for patients infected with COVID were often based on pulse oximeter readings (Valbuena, Merchant, &amp; Hough 2022). The problem is the device overestimates oxygen saturation in patients with darker skin, an issue which has been recognized for over thirty years (Valbuena, Merchant, &amp; Hough 2022). It would be as if a standard thermometer reported lower body temperatures for patients of colour, making it seem as if they did not have a fever when they actually did. In the case of COVID-19, Asians, Hispanics, and African Americans experienced inaccurately high readings of their oxygen levels (with African Americans and darker-skinned Hispanics having the highest), resulting in delays in treatment, hospital admission, and access to medications to treat COVID and contributing to higher severity of illness and higher death rates among these populations in comparison to whites (Fawzi Et al. 2022).<\/p>\n<p class=\"import-Normal\">Employment was also a factor in unequal exposure to and death from COVID-19 (Raifman, Skinner, &amp; Sojourner 2022), with many low-income workers making the choice (which, realistically, may not be a choice at all) to expose themselves to COVID in order to earn the funds necessary to purchase food, housing, and other necessities. Many such workers were then forced to miss work due to COVID infection. With only 35% of low-wage workers (as opposed to 95% of high-wage workers) having paid sick leave, this left many families struggling financially. Three years into the pandemic, low-wage workers continue to have the least access to COVID vaccines and boosters. The U.S. also lacks federal workplace-safety regulations with regard to vaccine and masking mandates that other nations enforce in times of high transmission, and it does not provide high-quality masks to its essential workers. Many occupations deemed essential by the CDC during the height of the pandemic\u2014such as health care, emergency services, meat packing, agricultural work, teaching, and jobs in the hospitality sector\u2014experienced higher rates of morbidity and mortality from COVID. Many of these fields disproportionately employ people of colour (McKinsey &amp; Company 2021). Given this, future policies that address the pandemic at a structural level\u2014for example, providing monetary assistance to people who work in environments with a high risk of infection, such as cleaning, nursing, transportation, retail, restaurant work, and factory work, so that they can remain at home\u2014may function more effectively to prevent transmission and curb future outbreaks (Arnot Et al. 2020).<\/p>\n<h2 class=\"import-Normal\">Food for Thought<\/h2>\n<p class=\"import-Normal\">This chapter focused primarily on health conditions prevalent in contemporary, industrialized societies that are due, in part, to the mismatch between our evolved biology and modern environments. These are the built environment and the social environment, which together form the obesogenic environment in which unhealthy behaviours are encouraged. This chapter will close by examining each of these in a college context.<\/p>\n<figure style=\"width: 275px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/opentextbooks.concordia.ca\/explorations\/wp-content\/uploads\/sites\/57\/2023\/08\/image2-3.jpg\" alt=\"Four individuals in a park.\" width=\"275\" height=\"183\" \/><figcaption class=\"wp-caption-text\">Figure 17.13: Students walking around a campus. Credit: <a href=\"https:\/\/www.maxpixel.net\/Row-Four-Man-Woman-People-Walking-Together-3755342\">Row four man woman people walking together 3755342<\/a> by <a href=\"https:\/\/www.maxpixel.net\/\">MaxPixel<\/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>Consider your campus from an evolutionary perspective. To what degree does the built environment lend itself to physical activity as part of daily life? Is your campus constructed in ways that promote driving at the expense of walking or biking? If driving is necessary, is parking available close to the buildings or do you need to walk a fair distance from the parking lot to your destination? Do the buildings have stairs or ramps or is it necessary to take the elevator? Is it possible to negotiate safely around campus on foot or by bike in all weather? After dark? How about the classrooms and computer labs? Do they have standing or treadmill desks? Does your class schedule encourage walking from building to building between classes, or are most courses in your major scheduled in the same location? Most college majors also lack a physical education requirement, leaving it up to students to incorporate exercise into already-challenging schedules (Figure 17.13).<\/p>\n<p class=\"import-Normal\">Sociocultural factors that contribute to obesity include food advertising, ubiquitous fast-food and junk food options, and social pressure to consume, all of which are present on college campuses. Although nutrition on campuses has improved in recent years, many students find eating healthy in the dining halls and dorms challenging (Plotnikoff Et al. 2015), and students who live off campus fare even worse (Small Et al<em>.<\/em> 2013). There are also parties and other social events, a normal part of college life, that involve unhealthy food and encourage behaviours like alcohol consumption and smoking. Give some thought to the social atmosphere on your campus and the ways it may contribute to obesity. My own freshman orientation involved a succession of pizza parties, ice cream socials, and barbecues, followed by late-night runs to the nearest fast-food outlet. The purpose of these events was to encourage people to make friends and feel comfortable living away from home, but the foods served were unhealthy, and there was social pressure to join in and be part of the group. Such activities set students up for the \u201cfreshman fifteen\u201d and then some. They also reinforce the idea that being social involves eating (and sometimes drinking and\/or smoking).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Sedentarism and inactivity are also built into the academics of college life. Digital technology is a significant contributor to obesity. Students use laptops and cell phones to take notes, complete their work outside of class, and access social media. There are also video games, virtual reality headsets, and streaming television and movies for entertainment. The built environment of college already necessitates that students sit in class for hours each day, then sit at computers to complete work outside of class. The social environment enabled by digital technology encourages sitting around for entertainment. It is telling that we call it \u201cbinge watching\u201d when we spend hours watching our favourite shows. Doing so often involves eating, as well as multiple exposures to food advertising embedded in the shows themselves. In these ways, college contributes to the development of obesity-causing behaviours that can have negative health ramifications long after college is over (Small Et al. 2013).<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">In the U.S., the greatest increase in obesity is among young adults aged 18\u201329 years, a significant percentage of whom are college students (Plotnikoff Et al. 2015). Analyses of college students\u2019 behaviour across semesters shows consumption of fruits and vegetables drops over time, as does the amount of physical activity, while consumption of sugar-sweetened beverages and fast-food goes up, leading to weight gain at nearly six times the rate of the general public (Small Et al. 2013). In response, many colleges and universities have instituted programs to encourage healthier eating and more physical activity among students (Plotnikoff Et al<em>.<\/em> 2015). It is important to emphasize that neither changes in diet or exercise are effective on their own.. A 2022 study of over 340,000 British participants demonstrated that physical activity and diet quality did not individually have an impact on cardiovascular disease or cancers (Ding Et al. 2022). That is, hitting the gym won\u2019t counteract the consequences of consuming high-calorie, fatty foods, and eating kale all day can\u2019t cancel out sedentary habits.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Just as no one fad diet is going to prove healthier than another, no one type of exercise is better than another. Anything that raises your heart rate and that you enjoy doing for at least an hour each day will work. Take advantage of opportunities to build exercise into everyday life. Take the stairs, park as far away from buildings as possible, ride a bike or walk instead of driving, and take walks between classes instead of sitting down and checking your phone. As far as diets go, eating a few less unhealthy calories each day, one less soda, no sugar in your coffee, or letting that last slice of pizza go to someone else, make a difference in the long run. Little changes add up to bigger ones. We cannot change our biology, but we can certainly change our habits.<\/p>\n<div class=\"textbox shaded\">\n<h2>Summary<\/h2>\n<p data-start=\"134\" data-end=\"760\">The health problems faced by modern humans can often be understood through the lens of evolution. As early <em>Homo sapiens<\/em> evolved in environments with a wide diversity of edible plants and animals, we developed complex nutritional requirements. However, with dramatic changes in our environments and daily lives, modern humans now encounter a variety of diet-related challenges. These include shifts in dentition and intestinal morphology, changes in food production and preparation, and altered patterns of physical activity. Together, these factors have contributed to changes in the leading causes of morbidity and mortality.<\/p>\n<p data-start=\"762\" data-end=\"1565\">These health consequences are often framed through the concept of epidemiological transitions, which are influenced by diet, physical activity, population density, and exposure to zoonotic diseases linked to agriculture. It is proposed that the first transition occurred when humans shifted from foraging to food production, leading to increased exposure to new diseases and dietary changes. The second transition followed the Industrial Revolution, when improvements in lifestyle and living conditions reduced infectious diseases but led to a rise in non-communicable diseases. The third and fourth transitions, still unfolding today, are marked by the spread of drug-resistant pathogens, the re-emergence of infectious diseases, and new health challenges driven by globalization, urbanization, and climate change.<\/p>\n<p data-start=\"1567\" data-end=\"1807\">Modern health issues such as obesity, cardiovascular disease, cancer, stress-related conditions, and syndemics illustrate the ongoing interaction between our evolved biology and the cultural and physical environments in which we live.<\/p>\n<h2 class=\"import-Normal\">Review Questions<\/h2>\n<ul>\n<li class=\"import-Normal\">Why do humans like foods that are \u201cbad\u201d for them? Describe the evolutionary underpinnings of our tastes for sugar, salt, and fat.<\/li>\n<li class=\"import-Normal\">How might understanding contemporary disease from an evolutionary perspective benefit medical practitioners in treating their patients?<\/li>\n<li class=\"import-Normal\">Several risk factors for conditions like heart disease, diabetes, and cancer are referred to as \u201clifestyle factors,\u201d implying these are behavioural choices people make that put them at risk. These include unhealthy eating, lack of physical activity, smoking, and alcohol consumption. To what degree is unhealthy behaviour structured by the physical and social environment? For example, how does being a college student influence your eating habits, physical activity patterns, smoking, and consumption of alcohol?<\/li>\n<li class=\"import-Normal\">Who benefits from the global obesity epidemic? Think about how the following industries and institutions might profit from it: The medical establishment? The fitness industry? The diet industry? Fashion? Pharmaceutical companies? Food manufacturers? Advertisers?<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<h2 class=\"__UNKNOWN__\">Key Terms<\/h2>\n<div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\"><strong>Body mass index (BMI)<\/strong>: A person\u2019s weight in kilograms divided by the square of their height in metres. This is the most widely used measure for identifying obesity. The formula using kilograms and metres (or centimetres) is: weight (kg) \/ [height (m)]<sup>2 <\/sup>. The formula using pounds and inches is: 703 x weight (lbs) \/ [height (in)]<sup>2 <\/sup>. Use of the BMI is controversial for several reasons, including that it does not take into account age, bone structure, muscle mass, fat distribution, or ethnic and racial differences in body type.<\/p>\n<p class=\"import-Normal\"><strong>Cancer<\/strong>: A collection of related diseases in which some of the body\u2019s cells begin to divide without stopping and spread into surrounding tissues.<\/p>\n<p class=\"import-Normal\"><strong>Cardiovascular disease (CVD)<\/strong>: A disease of the heart and blood vessels, often related to atherosclerosis. CVD is a condition in which a substance called plaque builds up in the walls of the arteries, the blood vessels that carry blood away from the heart, which compromises the flow of blood to the heart or brain.<\/p>\n<p class=\"import-Normal\"><strong>Central nervous system<\/strong>: The complex of nerve tissues stemming from the brain and spinal cord that controls the activities of the body.<\/p>\n<p class=\"import-Normal\"><strong>Circulatory (system)<\/strong>: The biological system that circulates blood around the body via the heart, arteries, and veins, delivering oxygen and nutrients to organs and cells and carrying waste products away.<\/p>\n<p class=\"import-Normal\"><strong>Diabetes mellitus<\/strong>: An endocrine disorder in which high glucose (blood sugar) levels occur over a prolonged period of time. Blood glucose is your body\u2019s main source of energy and comes from the food you eat. Insulin, a hormone made by the pancreas, helps glucose from food get into your cells to be used for energy. Sometimes your body does not make enough\u2014or any\u2014insulin (type 1 diabetes) or does not take up insulin well (type 2 diabetes). Glucose then stays in your blood and does not reach your cells.<\/p>\n<p class=\"import-Normal\"><strong>\u201cDouble burden\u201d<\/strong>: Refers to parts of the world in which there is a prevalence of chronic disease (e.g., cancer, heart disease) while, at the same time, there are also high rates of infectious disease due to poverty, malnutrition, poor sanitation, and lack of access to health care, often accompanied by high rates of maternal and child mortality.<\/p>\n<p class=\"import-Normal\"><strong>Ecological model<\/strong>: Ecological models of health and disease emphasize environmental and policy contexts of behaviour, while incorporating social and psychological influences, rather than focusing on individual behaviours. These models encompass multiple levels of influence and can lend themselves to more comprehensive health interventions.<\/p>\n<p class=\"import-Normal\"><strong>Emerging infectious diseases (EIDs)<\/strong>: Infections that have recently appeared within a population or those whose incidence or geographic range is rapidly increasing or threatens to increase in the near future. Examples include Covid-19, Ebola, Zika, SARS, and avian (bird) flu.<\/p>\n<p class=\"import-Normal\"><strong>Endocrine system<\/strong>: Those organs in the body whose primary function is the production of hormones.<\/p>\n<p class=\"import-Normal\"><strong>Epidemiological transition<\/strong>: A transformation in patterns of disease (morbidity) and death (mortality) among a population.<\/p>\n<p class=\"import-Normal\"><strong>Glycemic index (GI)<\/strong>: A system that ranks foods on a scale from 1 to 100 based on their effect on blood-sugar levels. Carbohydrates with a low GI value (55 or less) are more slowly digested and metabolized causing a lower, slower rise in blood glucose and insulin levels.<\/p>\n<p class=\"import-Normal\"><strong>Hypertension<\/strong>: High blood pressure. Blood pressure is the force exerted by the blood against the walls of the blood vessels. In a blood pressure reading, the top number (usually higher) refers to the systolic pressure, the amount of pressure in your arteries during the contraction of your heart muscle when your heart beats. The bottom number is the diastolic pressure when your heart muscle is resting between beats. Hypertension can lead to severe health complications and increases the risk of heart attack and stroke.<\/p>\n<p class=\"import-Normal\"><strong>Incidence<\/strong>: The rate at which new cases of a disease occur in a population over a given period of time.<\/p>\n<p class=\"import-Normal\"><strong>Insulin<\/strong>: A hormone produced in the pancreas that regulates the amount of glucose in the blood. Lack of insulin or the inability to absorb insulin causes diabetes.<\/p>\n<p class=\"import-Normal\"><strong>Metabolic syndrome<\/strong>: A cluster of conditions, including increased blood pressure, high blood sugar, excess body fat around the waist, and abnormal cholesterol levels that occur together, increasing the risk of heart disease, stroke, and diabetes. Lifestyle changes like losing weight, exercising regularly, and making dietary changes can help prevent or reverse metabolic syndrome.<\/p>\n<p class=\"import-Normal\"><strong>Menarche<\/strong>: The first occurrence of menstruation.<\/p>\n<p class=\"import-Normal\"><strong>Morbidity<\/strong>: The number of cases of disease per unit of population occurring over a unit of time.<\/p>\n<p class=\"import-Normal\"><strong>Mortality<\/strong>: The number of deaths attributable to a particular cause per unit of population over a unit of time.<\/p>\n<p class=\"import-Normal\"><strong>Noncommunicable diseases (NCDs)<\/strong>: Also known as chronic diseases, NCDs tend to be of long duration and are the result of a combination of genetic, physiological, environmental, and behaviour factors. The main types of NCDs are cardiovascular diseases (like heart attacks and stroke<strong>)<\/strong>, cancers, chronic respiratory diseases (such as chronic obstructive pulmonary disease and asthma), and diabetes.<\/p>\n<p class=\"import-Normal\"><strong>Obesity<\/strong>: A medical condition in which excess body fat has accumulated to the point that it has adverse effects on health. Although controversial due to its lack of ethnic and racial specificity, the most widely used measure for identifying obesity is the body mass index (BMI), a person\u2019s weight in kilograms divided by the square of their height in metres. A measure of 30 kg\/m<sup>2<\/sup> is considered obese and 25\u201329 kg\/m<sup>2<\/sup> is considered overweight. Distribution of body fat also matters. Fat in the abdominal region has a stronger association with type 2 diabetes and cardiovascular disease, meaning waist-to-hip ratio and waist circumference are also important indicators of obesity-related health risk.<\/p>\n<p class=\"import-Normal\"><strong>Obesogenic<\/strong>: Promoting excessive weight gain.<\/p>\n<p class=\"import-Normal\"><strong>Omnivorous<\/strong>: Able to eat and digest foods of both plant and animal origins.<\/p>\n<p class=\"import-Normal\"><strong>Osteoarthritis<\/strong>: Refers to the degeneration of joint cartilage and underlying bone, causing pain and stiffness. In the absence of previous injury, it is most common in modern populations from middle age onward.<\/p>\n<p class=\"import-Normal\"><strong>Prevalence<\/strong>: The proportion of individuals in a population who have a particular disease or condition at a given point in time.<\/p>\n<p class=\"import-Normal\"><strong>Sedentarism<\/strong>: A way of life characterized by much sitting and little physical activity.<\/p>\n<p class=\"import-Normal\"><strong>Sedentism<\/strong>: Living in groups settled permanently in one place.<\/p>\n<p class=\"import-Normal\"><strong>Stress response<\/strong>: A predictable response to any significant threat to homeostasis. The human stress response involves the <strong>Central Nervous System<\/strong> and the endocrine system acting together. Sudden and severe stress incites the \u201cflight or flight\u201d response from the autonomic nervous system in conjunction with hormones secreted by the adrenal and pituitary glands, increasing our heart rate and breathing and releasing glucose from the liver for quick energy.<\/p>\n<p class=\"import-Normal\"><strong>Stroke<\/strong>: A stroke occurs when a blood vessel leading to the brain is blocked or bursts, preventing that part of the brain from receiving blood and oxygen, leading to cell death.<\/p>\n<p class=\"import-Normal\"><strong>Syndemic<\/strong>: The aggregation (grouping together) of two or more diseases or health conditions in a population in which there is some level of harmful biological or behavioural interface that exacerbates the negative health effects of any or all of the diseases involved. Syndemics involve the adverse interaction of diseases of all types, including infections, chronic noncommunicable diseases, mental health problems, behavioural conditions, toxic exposure, and malnutrition.<\/p>\n<p class=\"import-Normal\"><strong>Tricep skinfold measurement<\/strong>: The triceps skinfold site is a common location used for the assessment of body fat using skinfold calipers. A section of skin on the posterior (back) surface of the arm that lays atop the tricep muscle is pinched between calipers. The resulting measurement is matched against a chart standardized for age and gender.<\/p>\n<p class=\"import-Normal\"><strong>\u201cTriple burden\u201d<\/strong>: A fourth epidemiological transition currently underway in which some parts of the globe are suffering from the \u201cdouble burden\u201d of infectious and chronic diseases combined with injuries and diseases related to intensifying globalization, urbanization, deforestation, and climate change.<\/p>\n<p class=\"import-Normal\"><strong>Vector-borne diseases<\/strong>: Human illnesses caused by parasites, viruses, and bacteria that are transmitted by mosquitoes, flies, ticks, mites, snails, and lice.<\/p>\n<p class=\"import-Normal\"><strong>Zoonoses<\/strong>: Diseases that can be transmitted from animals to humans.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration<strong><br \/>\n<\/strong><\/h2>\n<p>Lents, Nathan H. 2018. <em>Human Errors: A Panorama of Our Glitches, from Pointless Bones to <\/em><em>Broken Genes<\/em>. Boston: Houghton Mifflin Harcourt.<\/p>\n<p>Stearns, Stephen C., and Jacob C. Koella, eds. 2008. <em>Evolution in Health and Disease<\/em>. 2nd edition. United Kingdom: Oxford University Press.<\/p>\n<p>Zuk, Marlene. 2013. <em>Paleofantasy: What Evolution Really Tells Us about Sex, Diet, and <\/em><em>How We Live<\/em>. New York: W. W. Norton &amp; Company.<\/p>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Abid, Zaynah, Amanda J. Cross, and Rashmi Sinha. 2014. \u201cMeat, Dairy, and Cancer.\u201d <em>The American Journal of Clinical Nutrition<\/em> 100 (S1): 386S\u2013393S.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Akil, Luma, and H. Anwar Ahmad. 2011. \u201cRelationships between Obesity and Cardiovascular Diseases in Four Southern States and Colorado.\u201d <em>Journal of Health Care for the Poor and Underserved<\/em> 22 (S4): 61\u201372.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Ali, Aus Tariq. 2014. \u201cReproductive Factors and the Risk of Endometrial Cancers.\u201d <em>International Journal of Gynecological Cancer<\/em> 24 (3): 384\u2013393.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">American Heart Association. 2018. \u201cHeart Disease and Stroke Statistics-2018 Update: A Report.\u201d <em>Circulation 137<\/em> (12). Accessed April 7, 2023. https:\/\/www.ahajournals.org\/doi\/10.1161\/cir.0000000000000558.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">American Psychiatric Association (APA). 2013. <em>Diagnostic and Statistical Manual of Mental Disorder<\/em>. 5th Edition: DSM-5. Washington, DC: APA.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Armelagos, George J. 1990. \u201cHealth and Disease in Prehistoric Populations in Transition.\u201d <em>Disease in Populations in Transition: Anthropological and Epidemiological Perspectives,<\/em> edited by George J. Armelagos and Alan C. Swedland, 127\u2013144. New York: Bergin &amp; Garvey.<\/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 and Medicine 61<\/em> (4): 755-765.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0.495002746582031pt;margin-right: 0pt;text-indent: 0pt\">Arnot, Megan, Eva Brandl, O. L. K. Campbell, Yuan Chen, Mark Dyble, Emily H. Emmott, et al. 2020. <em>Evolution, Medicine, and Public Health<\/em> 2020 (1): 264\u2013278. https:\/\/doi.org\/10.1093\/emph\/eoaa038.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Asher, Claire. 2017. \u201cIllegal Bushmeat Trade Threatens Human Health and Great Apes.\u201d <em>Mongabay<\/em>, April 6. Accessed April 4, 2023. https:\/\/news.mongabay.com\/2017\/04\/illegal-bushmeat-trade-threatens-human-health-and-great-apes\/.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Baker, W. A., G. A. Hitman, K. Hawrami, M .I. McCarthy, A. Riikonen, E. Tuomilehto-Wolf, A. Nissinen, et al. 1994. \u201cApolipoprotein D Gene Polymorphism: A New Genetic Marker for Type 2 Diabetic Subjects in Nauru and South India.\u201d <em>Diabetic Medicine<\/em> 11 (10): 947\u2013952.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Baltic, Milan Z., and Marija Boskovic. 2015. \u201cWhen Man Met Meat: Meat in Human Nutrition from Ancient Times Till Today.\u201d <em>Procedia Food Science 5<\/em>: 6- 9.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Blue Cross Blue Shield Association (BCBSA). 2017. \u201cDiabetes and the Commercially Insured U.S. Population.\u201d <em>The Health of America Report<\/em>, August 1. Accessed April 4, 2023. https:\/\/www.bcbs.com\/the-health-of-america\/reports\/diabetes-and-commercially-insured-us-population.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Bogin, Barry. 1991. \u201cThe Evolution of Human Nutrition.\u201d In <em>The Anthropology of Medicine: From Culture to Method<\/em>, edited by Lola Romanucci-Ross, Daniel E. Moerman, and Laurence R. Tancredi, 158\u2013195. New York: Bergin &amp; Garvey.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Bouchard, Claude. 2007. \u201cThe Biological Predisposition to Obesity: Beyond the Thrifty Genotype Scenario.\u201d <em>International Journal of Obesity<\/em> 31 (9): 1337\u20131339.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Buchman, Aron S., Lei Yu, Robert S. Wilson, Andrew Lim, Robert J. Dawe, Chris Gaiteri, Sue E. Leurgans, Julie A. Schneider, and David A. Bennett. 2019. \u201cPhysical Activity, Common Brain Pathologies, and Cognition in Community-Dwelling Older Adults.\u201d <em>Neurology<\/em> 98 (2). https:\/\/doi.org\/10.1212\/WNL.0000000000006954.<\/p>\n<p class=\"import-Normal\" style=\"background-color: transparent;text-align: left;margin-left: 0pt;margin-right: 0pt;text-indent: 0pt\">Centers for Disease Control and Prevention (CDC). 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Wood. 2016. \u201cPhysical Activity Patterns and Biomarkers of Cardiovascular Disease Risk in Hunter-Gatherers.\u201d <em>American Journal of Human Biology<\/em> 29 (2): e22919. https:\/\/doi.org\/10.1002\/ajhb.22919.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Raifman, Julia, Alexander Skinner, and Aaron Sojourner. 2022. \u201cThe Unequal Toll of COVID-19 On Workers.\u201d Working Economics Blog (of the Economic Policy Institute), February 7. Accessed April 4, 2023. <a class=\"rId84\" href=\"https:\/\/www.epi.org\/blog\/the-unequal-toll-of-covid-19-on-workers\/\">https:\/\/www.epi.org\/blog\/the-unequal-toll-of-covid-19-on-workers\/<\/a>.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Randolph-Quinney, Patrick S., Scott A. Williams, Maryna Stein, Mark R. Meyer, Jacqueline S. Smilg, Steven E. Churchill, Edward J. 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Hoboken, NJ: Wiley-Blackwell.<\/p>\n<p class=\"import-Normal\" style=\"margin-left: 0pt;text-indent: 0pt\">Zuckerman, Molly Kathleen, Kristin Nicole Harper, Ronald Barrett, and George John Armelagos. 2014. \u201cThe Evolution of Disease: Anthropological Perspectives on Epidemiologic Transitions.\u201d Special issue, \u201cEpidemiological Transitions: Beyond Omran\u2019s Theory,\u201d <em>Global Health Action<\/em> 7 (1): 23303. https:\/\/doi.org\/10.3402\/gha.v7.23303.<\/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_1683_1702\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1702\"><div tabindex=\"-1\"><p>Dr. Salinda Hess adopted Explorations: An Open Invitation to Biological Anthropology for ANTH 203: Culture and Biology at Concordia University in 2023. To reflect the ongoing changes and developments within the field of biological anthropology and to integrate Canadian perspectives, her syllabus included an open assignment titled \u201cGroup Participation in Open Pedagogy Textbook Project.\u201d The resulting student chapter-level feedback from this interactive assignment led to revisions, updates, corrections, and\/or added materials.<\/p>\n<p><strong>Lola Leus<\/strong> helped review and select student projects for inclusion in the textbook. She worked on integrating the selected and edited submissions into their designated chapters, ensuring that the additions maintained the flow, clarity, and consistency of the existing text. She also reviewed and incorporated recommended revisions from the first edition, helping finalize changes intended to improve the textbook\u2019s relevance and educational value. These revisions and student contributions were incorporated into the adapted and annotated second edition and further integrated into the third edition.<\/p>\n<p><strong>Adam Garcia<\/strong> played an editorial role and a technical Pressbooks role. He helped select insights from student assignments, checked citations, and integrated revisions in the customized text, which is now the third edition.<\/p>\n<table aria-labelledby=\"table-title\">\n<caption id=\"table-title\">Student Contributors by Chapter<\/caption>\n<thead>\n<tr>\n<th scope=\"col\">Chapter Title<\/th>\n<th scope=\"col\">Student Contributors<\/th>\n<\/tr>\n<\/thead>\n<tbody>\n<tr>\n<td>3. Social and Biopolitical Dimensions of Evolutionary Thinking<\/td>\n<td>Daphn\u00e9e-Tiffany Kirouac Millan, Davina Paradis, Jung Jin Kim, and Nathan Dennis<\/td>\n<\/tr>\n<tr>\n<td>4. Molecular Biology and Genetics<\/td>\n<td>Emma Costa, Shima Gahima, Will Lefebvre, Audrey Ch\u00e9kina\u00ebl<\/td>\n<\/tr>\n<tr>\n<td>5. Forces of Evolution<\/td>\n<td>Corin Laberge, Hazel Moorcroft, Isabella Michel, Julian J. Pantoja Quiroz<\/td>\n<\/tr>\n<tr>\n<td>8. Stones and Bones: Studying the Fossil Record<\/td>\n<td>Catherine Belec, Maria Papadakis, Camille Senior and Nadjat Baril<\/td>\n<\/tr>\n<tr>\n<td>10. Early Hominins<\/td>\n<td>Angela Durastanti, Bryce Muller, Gabriel Barr, Maisie Babbington-Bolduc<\/td>\n<\/tr>\n<tr>\n<td>12. Archaic Homo<\/td>\n<td>Peyton Dagg, Bryana Henry, and Anoriel Jacques<\/td>\n<\/tr>\n<tr>\n<td>16. Forensic Anthropology<\/td>\n<td>Amelia Roberts, Elyse Racicot, Emmanuelle Hunter<\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1683_1704\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1704\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<div class=\"textbox\">\n<h6>Student contributors for this chapter from Salinda Hess' ANTH 203: Daphn\u00e9e-Tiffany Kirouac Millan, Davina Paradis, Jung Jin Kim, and Nathan Dennis<\/h6>\n<p>Learn more about <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/about-the-authors\/\">authors and editors<\/a>,<a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/student-contributors\/\"> Hess' student contributions<\/a>, and <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/versioning-history\/\">versioning history<\/a>.<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\">Learning Objectives<\/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 flavour 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 flavours 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. Lewontin, Rose, and Kamin (2017) argue that the same is true for humans and other organisms.<\/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 acknowledgement 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 3.1: 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 3.1). 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 &amp; 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 &amp; 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 3.2). 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, &amp; 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 3.2: 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 neighbouring 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 3.3).<\/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 3.3: 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 analyse 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_1683_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 favourable traits. Both Waddington and Gould resisted contemporary intellectual paradigms that favoured 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 labour, 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 4). 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 4). 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 4).<\/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 behavioural variation and culture. That is, behavioural 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 proposed 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, behavioural, 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<div class=\"textbox\" style=\"background: var(--lightblue)\">\n<h2>Dig Deeper: Moving Beyond Genetic Determinism<\/h2>\n<p>Contemporary evolutionary biology and anthropology increasingly emphasize that genes operate within dynamic regulatory networks rather than acting as isolated determinants. As <a href=\"https:\/\/www.zotero.org\/google-docs\/?zoqFM1\">Carroll (2005)<\/a> and <a href=\"https:\/\/www.zotero.org\/google-docs\/?C6NEFg\">Wray (2007)<\/a> demonstrate, evolutionary change often arises not from mutations in structural genes but in their regulation\u2014the timing, intensity, and location of gene expression. Such regulatory evolution can explain major anatomical and physiological innovations without invoking large genetic divergences. This view reframes evolution as an outcome of organizational complexity where genetic, developmental, and environmental processes intersect. This systems-level understanding also resonates with anthropological frameworks of biocultural embodiment, which recognize that social and ecological experiences can become biologically inscribed in the body. <a href=\"https:\/\/www.zotero.org\/google-docs\/?AROEum\">Meaney\u2019s (2001)<\/a>\u00a0 foundational epigenetic research focuses on maternal care in rats, presenting how nurturing behaviour modifies the expression of stress-response genes. This biological effect can persist into subsequent generations.<\/p>\n<p>Recent human studies continue to expand this insight. <a href=\"https:\/\/www.zotero.org\/google-docs\/?r3ZGNw\">Goldman &amp; Sterner (2023)<\/a> demonstrate how environmental exposures, inequality, and psychological stress influence the pace of biological aging, showing epigenetic modifications reflect the lived conditions of bodies over time. In Canada, this relationship between environment, history, and biology has profound implications. A 2023 scoping review on Canadian Indigenous populations and the epigenetic effects of intergenerational trauma <a href=\"https:\/\/www.zotero.org\/google-docs\/?NEGUdK\">(Schafte &amp; Bruna, 2023)<\/a> documents measurable biological patterns associated with colonial violence, displacement, and systemic inequity. By dissecting the obesity patterns in the Indigenous youth populations, the researchers present a clear connection between the parents who attended residential schools and biological health issues in their children years later. This holistic understanding of epigenetics shows an \u201cembodied transmission of trauma and ill health across generations\u201d (2023, p.9), underscoring that the effects of colonialism are not merely social but are biologically embodied, carried forward through mechanisms of gene regulation and stress physiology.<\/p>\n<p>Understanding heredity as a process of interaction and regulation rather than genetic determinism opens the door to rethinking evolution as a flexible, context-driven phenomenon. Just as social experiences and ecological conditions can shape patterns of gene expression, environmental pressures can also influence the structure and behaviour of genomes across generations. This broader view of evolutionary change highlights the importance of considering mechanisms that fall outside of traditional, gradualist models.<\/p>\n<\/div>\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 3.4). 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 such as 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 3.4: 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 &amp; McKie 1997), <em>The Human Blueprint <\/em>(Shapiro 1991), and <em>The Code of Codes<\/em> (Kevles &amp; 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<\/div>\n<p>&nbsp;<\/p>\n<div class=\"textbox\" style=\"background: var(--lightblue)\">\n<h2><strong>Special Topic: Oversexing the Gendered Body <\/strong><\/h2>\n<p>While rapid mitochondrial evolution underscores the biological flexibility of organisms in response to environmental pressures, evolutionary theory is also shaped by another set of forces: cultural assumptions and social norms. Nowhere is this more visible than in scientific interpretations of sex and gender. Many modern gender roles stem from assumptions about sex differences that have accumulated throughout human history. While these roles may appear to be fixed stereotypes or biologically predetermined, they can be deconstructed by examining the processes of sexual selection through queer and feminist theoretical frameworks. Applying these lenses to evolutionary concepts allows for a deeper understanding of how cultural ideologies, particularly those surrounding gender and sexuality, shape interpretations of biological processes.<\/p>\n<p>Darwin first introduced the concept of sexual selection in The Descent of Man (1871) to explain how males and females may have developed different traits that would be detrimental to the species\u2019 overall survival <a href=\"https:\/\/www.zotero.org\/google-docs\/?GVyarx\">(Vicedo, 2025)<\/a>. Unlike natural selection which is \u201cselection by death,\u201d sexual selection represents death by selection <a href=\"https:\/\/www.zotero.org\/google-docs\/?G5vjwZ\">(Gayon, 2010)<\/a>. Darwin argued that males typically compete intrasexually for female attention, and that females exercise choice based on attractiveness or vigor, proving their fitness. However, when reframed through feminist theory, the amount of agency Darwin ascribed to females doesn\u2019t reflect the societal assumptions surrounding gender roles in his era. Charlotte Perkins Gilman in her publication Women and Economics, argued that by the 1960s, men increasingly relied on social dominance over women rather than competition with other men (Vicedo, 2025). This dynamic required women to continually enhance their sexual appeal in exchange for economic security, a system she coined the \u201csexuo-economic relationship\u201d (2025, p.5). This framework reveals the societal power imbalance between men and women, and how women are the ones sexualizing themselves and competing for partners, not men. Such processes would lead to the modern oversexualization of women.<\/p>\n<p>Oversexualization, a cultural ideology that prioritizes sexual appeal over autonomy and well-being, further complicates interpretations of sexual selection. Brassard and company (2018) define oversexualization through four components: valuing people solely for their sexual appeal, societal norms of equating attractiveness with sexiness, sexual objectification, and the inappropriate imposition of sexuality (Brassard Et al., 2018, p.16-17). When oversexualization is observed within a population, it may signal that the pressures of sexual selection have intensified relative to that of natural selection, creating \u201cexcessive sex difference\" (Vicedo,<a href=\"https:\/\/www.zotero.org\/google-docs\/?a1BV2F\"> 2025)<\/a>. While many aspects of Gilman's arguments do not directly apply to contemporary gender dynamics, stereotypes rooted in historical gender expectations continue to shape women's experiences in the workforce and broader society (2025). Understanding sexual selection as a culturally mediated process, rather than as a simple competition amongst males, offers a more nuanced picture of how gender ideologies influence biological narratives. This intersection of culture and biology is crucial for studying gender roles, queer relationships, and sexual diversity across societies and time periods.<\/p>\n<\/div>\n<p>&nbsp;<\/p>\n<div class=\"__UNKNOWN__\">\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 suite<\/p>\n<figure style=\"width: 279px\" class=\"wp-caption alignright\"><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 3.5: 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\">d 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. Consid<\/p>\n<p class=\"import-Normal\">er 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<p>&nbsp;<\/p>\n<div class=\"textbox\" style=\"background: var(--lightblue)\">\n<h2>Dig Deeper: Rapid Mitochondrial Evolution in Stingless Bees<\/h2>\n<p>A striking example of this interactional evolutionary change comes from recent research on stingless bees. When observing the mitochondrial genome (mitogenome) in two Australian stringless bees in the genus Tetragonula\u2014T. carbonaria and T. hockingsi\u2014they exhibit a rare\u00a0 duplication of the entire mitogenome and show rapid divergence from other members of their species <a href=\"https:\/\/www.zotero.org\/google-docs\/?yfSvM1\">(Fran\u00e7oso et al., 2023)<\/a>. This accelerated evolution is hypothesized to result from factors such as low effective population, founder effects, and genome duplication triggered by environmental stressors. This phenomenon echoes the earlier work by Conrad H. Waddington (1956) mentioned in this chapter, whose experiments exposing fruit fly embryos to ether induced the development of additional wings and thoraces, changes that later became heritable under stable conditions <a href=\"https:\/\/www.zotero.org\/google-docs\/?FdVXeR\">(Shook et al., 2023b)<\/a>. Both cases highlight how organisms can respond to intense environmental pressures through dramatic developmental and genetic shifts.<\/p>\n<p>The mitochondria genetics influence the energy synthesis of the cells and in most animals, the mitogenome remains relatively stable <a href=\"https:\/\/www.zotero.org\/google-docs\/?Lw5Lmk\">(Shook et al., 2023a)<\/a>; however, Tetragonula species appear to possess an unusual capacity for rapid sequence rearrangement and complete genome duplication, suggesting that their mitogenomes play an important adaptive role. Comparing these genomes with other species such as Lepidotrigona\u2014which shows rearrangements but no duplication\u2014 provides a unique opportunity to examine how different lineages respond to similar ecological pressures. <a href=\"https:\/\/www.zotero.org\/google-docs\/?VxrGpj\">Fran\u00e7oso et al. (2023)<\/a> found that Tetragonula mitogenomes form amphimeric circular structures in which two complete genomes are joined head-to-tail, an extremely rare configuration. These arrangements, including inversions and translocations of gene blocks such as ND6, CytB, ND1, and several rRNA and tRNA genes, are far less common in other bee genera. This pattern supports the idea proposed by <a href=\"https:\/\/www.zotero.org\/google-docs\/?V2eJWI\">Gould &amp; Eldredge (1977)<\/a> that species are fundamentally unstable entities subject to bursts of rapid change in response to environmental pressures, rather than progressing along a single linear pathway. It is important to note that not all species within the genus exhibit the same degree or type of genomic flexibility. While T. carbonaria and T. hockingsi show full mitogenome duplications, the aforementioned Lepidotrigona species show only partial rearrangements despite facing similar environmental conditions. This variation challenges deterministic assumptions that evolution necessarily moves species toward optimal forms. Instead, it illustrates that evolution often involves trial-and-error shifts shaped by constraint, chance, and ecological stress.<\/p>\n<p>Although further research is needed to determine precisely what triggers such rapid genomic events, the evidence demonstrates that mitochondria play an active role in shaping evolutionary pathways. These findings complicate traditional gradualist models and highlight the importance of examining molecular responses to environmental pressures.<\/p>\n<\/div>\n<p>&nbsp;<\/p>\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 3.5). 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 3.6: 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 behaviour could be mapped to distinct parts of the brain and physically observed (see Figure 3.6). 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 3.7: 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 behaviour 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 3.7). 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, behaviourally, 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 favour 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_1683_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 labour, 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 behaviours are also highly adaptive, so what about our most familiar social behaviours? 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 harboured 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\" style=\"background: var(--lightblue)\">\n<p class=\"import-Normal\"><strong style=\"font-family: 'Cormorant Garamond', serif;font-size: 1.602em\">Evolution of the Anthropocene <\/strong><\/p>\n<figure style=\"width: 379px\" class=\"wp-caption alignright\"><img class=\"\" src=\"https:\/\/upload.wikimedia.org\/wikipedia\/commons\/thumb\/8\/8f\/Absetzterseite_des_Tagebaus_Inden_2002.jpg\/500px-Absetzterseite_des_Tagebaus_Inden_2002.jpg\" alt=\"File:Absetzterseite des Tagebaus Inden 2002.jpg\" width=\"379\" height=\"200\" \/><figcaption class=\"wp-caption-text\">Figure 3.8: View of the overburden dumping side of the Inden open-pit lignite mine in the Rhineland, Germany, showing layers of excavated earth used to reconstruct the landscape. Credit: <em data-start=\"249\" data-end=\"289\">Absetzterseite des Tagebaus Inden 2002<\/em> by Rhetos is dedicated to the public domain under the <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/deed.en\">Creative Commons CC0 1.0 Universal Public Domain Dedication. <\/a><\/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<\/div>\n<\/div>\n<div class=\"__UNKNOWN__\">\n<h2 class=\"import-Normal\"><span style=\"background-color: #ffffff\">Summary<\/span><\/h2>\n<p class=\"import-Normal\"><span style=\"background-color: #ffffff\">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: #ffffff\">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 favourable 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\">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\">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>. 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Social and Biopolitical Dimensions of Evolutionary Thinking. https:\/\/opentextbooks.concordia.ca\/explorationsversiontwo\/chapter\/social-and-bi opolitical-dimensions-of-evolutionary-thinking\/<\/p>\n<p class=\"import-Normal\">Spencer, Herbert. 1864. <em>Principles of Biology.<\/em> London: Williams and Norgate.<\/p>\n<p>UNESCO. (2024).<em> The Anthropocene<\/em>. International Union of Geological Sciences. https:\/\/www.iugs.org\/_files\/ugd\/f1fc07_40d1a7ed58de458c9f8f24de5e739663.pdf?index=true<\/p>\n<p>Vicedo, M. (2025). Charlotte Perkins Gilman: A Pragmatist Framework for Constructing a New Humanhood. Journal of the History of the Behavioral Sciences, 61(4), e70041. MEDLINE. https:\/\/doi.org\/10.1002\/jhbs.70041<\/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>Wray, G. A. (2007). The evolutionary significance of cis-regulatory mutations. Nature Reviews. Genetics, 8(3), 206\u2013216. MEDLINE.<\/p>\n<p>Waddington CH. Genetic assimilation of the bithorax phenotype. Evolution. 1956;10:1\u201313. doi: 10.1111\/j.1558-5646.1956.tb02824.x.<\/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>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><template id=\"term_1683_1705\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1705\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<div class=\"textbox\">\n<h6>Student contributors for this chapter from Salinda Hess' ANTH 203: <em>Emma Costa, Shima Gahima, Will Lefebvre, Audrey Ch\u00e9kina\u00ebl. <\/em><\/h6>\n<p>Learn more about <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/about-the-authors\/\">authors and editors<\/a>,<a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/student-contributors\/\"> Hess' student contributions<\/a>, and <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/versioning-history\/\">versioning history<\/a>.<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\">Learning Objectives<\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li class=\"import-Normal\">Outline a 21st-century perspective of the Modern Synthesis.<\/li>\n<li class=\"import-Normal\">Define populations and population genetics as well as the methods used to study them.<\/li>\n<li class=\"import-Normal\">Identify the forces of evolution and become familiar with examples of each.<\/li>\n<li class=\"import-Normal\">Discuss the evolutionary significance of mutation, genetic drift, gene flow, and natural selection.<\/li>\n<li class=\"import-Normal\">Explain how allele frequencies can be used to study evolution as it happens.<\/li>\n<li class=\"import-Normal\">Contrast micro- and macroevolution.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p>It\u2019s hard for us, with our typical human life spans of less than 100 years, to imagine all the way back, 3.8 billion years ago, to the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1442\">origins of life<\/a><\/strong>. Scientists still study and debate how life came into being and whether it originated on Earth or in some other region of the universe (including some scientists who believe that studying evolution can reveal the complex processes that were set in motion by God or a higher power). What we do know is that a living single-celled organism was present on Earth during the early stages of our planet\u2019s existence. This organism had the potential to reproduce by making copies of itself, just like bacteria, many amoebae, and our own living cells today. In fact, with modern technologies, we can now trace genetic lineages, or <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1290\">phylogenies<\/a><\/strong>, and determine the relationships between all of today\u2019s living organisms\u2014eukaryotes (animals, plants, fungi, etc.), archaea, and bacteria\u2014on the branches of the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1444\">phylogenetic tree of life<\/a><\/strong> (Figure 5.1).<\/p>\n<figure style=\"width: 675px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2023\/02\/image1-1.png\" alt=\"Branches lead off of a single celled universal ancestor to images of bacteria, archaea, and eukarya (represented by a mouse, mushroom, and fern, among others).\" width=\"675\" height=\"475\" \/><figcaption class=\"wp-caption-text\">Figure 5.1: Phylogenetic tree of life illustrating probable relationships between the single-celled Last Universal Common Ancestor (LUCA) and select examples of bacteria, archaea, and eukaryotes. Major evolutionary developments, including independent evolution of multicellularity, photosynthesis, and respiration, are indicated along the branches. <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 class=\"rId11\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Cladograma_dos_Dom%C3%ADnios_e_Reinos.png\">Cladograma dos Dominios e Reinos<\/a> by <a class=\"rId12\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:MarceloTeles\">MarceloTeles<\/a> has been modified (English labels replace Portuguese) and is under a <a class=\"rId13\" 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\">Looking at the common sequences in modern genomes, we can even make educated guesses about the likely genetic sequence of the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1446\"><strong>Last Universal Common Ancestor (LUCA)<\/strong><\/a> of all living things. Through a wondrous series of mechanisms and events over nearly four billion years, that ancient single-celled organism gave rise to the rich diversity of species that fill the lands, seas, and skies of our planet. This chapter explores the mechanisms by which that amazing transformation occurred and considers some of the crucial scientific experiments that shaped our current understanding of the evolutionary process.<\/p>\n<h2 class=\"import-Normal\">Population Genetics<\/h2>\n<h3 class=\"import-Normal\"><strong>Defining Populations and the Variations <\/strong><strong>w<\/strong><strong>ithin Them<\/strong><\/h3>\n<p class=\"import-Normal\">One of the major breakthroughs in understanding the mechanisms of evolutionary change came with the realization that evolution takes place at the level of populations, not within individuals. In the biological sciences, a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1454\"><strong>p<\/strong><strong>opulation<\/strong><\/a> is defined as a group of individuals of the same <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1456\"><strong>species<\/strong><\/a> who are geographically near enough to one another that they can breed and produce new generations of individuals.<\/p>\n<p class=\"import-Normal\">For the purpose of studying evolution, we recognize populations by their even smaller units: genes. Remember, a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1458\"><strong>gene<\/strong><\/a> is the basic unit of information that encodes the proteins needed to grow and function as a living organism. Each gene can have multiple <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_738\">alleles<\/a><\/strong>, or variants\u2014each of which may produce a slightly different protein. Each individual, for genetic inheritance purposes, carries a collection of genes that can be passed down to future generations. For this reason, in population genetics, we think of populations as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1462\"><strong>gene pools<\/strong><\/a>, which refers to the entire collection of genetic material in a breeding community that can be passed on from one generation to the next.<\/p>\n<p class=\"import-Normal\">For genes carried on our human chromosomes (our nuclear DNA), we inherit two copies of each, one from each parent. This means we may carry two of the same alleles (a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1464\"><strong>homozygous genotype<\/strong><\/a>) or two different alleles (a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1466\"><strong>heterozygous<\/strong> <strong>genotype<\/strong><\/a>) for each nuclear gene.<\/p>\n<h3 class=\"import-Normal\"><strong>Defining Evolution <\/strong><\/h3>\n<p class=\"import-Normal\">In order to understand evolution, it\u2019s crucial to remember that evolution is always studied at the population level. Also, if a population were to stay exactly the same from one generation to the next, it would not be evolving. So evolution requires both a population of breeding individuals and some kind of a genetic change occurring within it. Thus, the simple definition of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1468\"><strong>evolution<\/strong><\/a> is a change in the allele frequencies in a population over time. What do we mean by allele frequencies? <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1470\"><strong>Allele frequencies<\/strong> <\/a>refer to the ratio, or percentage, of one allele (one variant of a gene) compared to the other alleles for that gene within the study population (Figure 5.2). By contrast, <strong>genotype frequencies<\/strong> are the ratios or percentages of the different homozygous and heterozygous genotypes in the population. Because we carry two alleles per <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_736\">genotype<\/a><\/strong>, the total count of alleles in a population will usually be exactly double the total count of genotypes in the same population (with the exception being rare cases in which an individual carries a different number of chromosomes than the typical two; e.g., Down syndrome results when a child carries three copies of Chromosome 21).<\/p>\n<figure style=\"width: 652px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image2.jpg\" alt=\"Genotypes are represented as combinations of alleles and allele frequencies.\" width=\"652\" height=\"883\" \/><figcaption class=\"wp-caption-text\">Figure 5.2: Population evolution can be measured by allele frequency changes. This diagram illustrates the differences between genotype frequencies and allele frequencies, as well as how they can be measured in a population of snapdragon flowers. The lower portion of the diagram also depicts how evolution is recognized as allele frequencies change in a population over time. <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: Population evolution original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Katie Nelson and Beth Shook is a collective work under a <a class=\"rId15\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\">CC BY-NC 4.0 License<\/a>. [Includes <a class=\"rId16\" href=\"https:\/\/pixabay.com\/vectors\/snapdragon-flower-pink-lilac-plant-146850\/\">Snapdragon-flower-pink-lilac<\/a> by <a class=\"rId17\" href=\"https:\/\/pixabay.com\/users\/openclipart-vectors-30363\/\">OpenClipart-Vectors<\/a>, <a class=\"rId18\" href=\"https:\/\/creativecommons.org\/share-your-work\/public-domain\/cc0\/\">public domain (CC0)<\/a> under a <a class=\"rId19\" href=\"https:\/\/pixabay.com\/service\/terms\/\">Pixabay License<\/a>.]<\/figcaption><\/figure>\n<h2 class=\"import-Normal\">The Forces of Evolution<\/h2>\n<p class=\"import-Normal\">Today, we recognize that evolution takes place through a combination of mechanisms: mutation, genetic drift, gene flow, and natural selection. These mechanisms are called the \u201cforces of evolution\u201d; together they account for all the genotypic variation observed in the world today. Keep in mind that each of these forces was first defined and then tested\u2014and retested\u2014through the experimental work of the many scientists who contributed to the Modern Synthesis.<\/p>\n<h3 class=\"import-Normal\"><strong>Mutation<\/strong><\/h3>\n<p class=\"import-Normal\">The first force of evolution we will discuss is mutation, and for good reason: mutation is the original source of all the genetic variation found in every living thing. Imagine all the way back in time to the very first single-celled organism, floating in Earth\u2019s primordial sea. Based on what we observe in simple, single-celled organisms today, that organism probably spent its lifetime absorbing nutrients and dividing to produce cloned copies of itself. While the numbers of individuals in that population would have grown (as long as the environment was favourable), nothing would have changed in that perfectly cloned population. There would not have been variety among the individuals. It was only through a copying error\u2014the introduction of a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1474\"><strong>mutation<\/strong><\/a>, or change, into the genetic code\u2014that new alleles were introduced into the population.<br style=\"clear: both\" \/><br style=\"clear: both\" \/>After many generations have passed in our primordial population, mutations have created distinct chromosomes. The cells are now amoeba-like, larger than many of their tiny bacterial neighbours, who have long since become their favourite source of nutrients. Without mutation to create this diversity, all living things would still be identical to LUCA, our universal ancestor (Figure 5.3).<\/p>\n<figure style=\"width: 663px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image3-2.png\" alt=\"Universal Ancestor linked to the Eukarya branch.\" width=\"663\" height=\"338\" \/><figcaption class=\"wp-caption-text\">Figure 5.3: Key mutational differences between Last Universal Common Ancestor and an amoeba-like primordial cell. <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<strong>: <\/strong>Key differences between LUCA and a primordial cell original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Andrea J. Alveshere is a collective work under a <a class=\"rId21\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Includes <a class=\"rId22\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Cladograma_dos_Dom%C3%ADnios_e_Reinos.png\">Cladograma dos Dominios e Reinos<\/a> by <a class=\"rId23\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:MarceloTeles\">MarceloTeles<\/a> (cropped, labels and colour changed), <a class=\"rId24\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0<\/a><a class=\"rId25\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">; <\/a><a class=\"rId26\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Amoeba_proteus_TK-UT.svg\">Amoeba Proteus TK-UT<\/a> by <a class=\"rId27\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Nefronus\">Tom\u00e1\u0161 Kebert<\/a> and <a class=\"rId28\" href=\"https:\/\/www.umimeto.org\/\">umimeto.org<\/a> (cropped and colour changed), <a class=\"rId29\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0<\/a>.]<\/figcaption><\/figure>\n<p class=\"import-Normal\">When we think of genetic mutation, we often first think of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1476\">deleterious mutations<\/a><\/strong>\u2014the ones associated with negative effects such as the beginnings of cancers or heritable disorders. The fact is, though, that every genetic adaptation that has helped our ancestors survive since the dawn of life is directly due to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1478\">beneficial mutations<\/a><\/strong>\u2014changes in the DNA that provided some sort of advantage to a given population at a particular moment in time. For example, a beneficial mutation allowed chihuahuas and other tropical-adapted dog breeds to have much thinner fur coats than their cold-adapted cousins the northern wolves, malamutes, and huskies.<\/p>\n<figure style=\"width: 320px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-1-1.png\" alt=\"UV radiation damages nucleotides in DNA.\" width=\"320\" height=\"248\" \/><figcaption class=\"wp-caption-text\">Figure 5.4: A crosslinking mutation in which a UV photon induces a bond between two thymine bases. <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<strong>: <\/strong><a class=\"rId31\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">UV-induced Thymine dimer mutation (Figure 4.6)<\/a> original to <a class=\"rId32\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a class=\"rId33\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Every one of us has genetic mutations. Yes, even you. The DNA in some of your cells today differs from the original DNA that you inherited when you were a tiny, fertilized egg. Mutations occur all the time in the cells of our skin and other organs, due to chemical changes in the nucleotides. Exposure to the UV radiation in sunlight is one common cause of skin mutations. Interaction with UV light causes <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1480\"><strong>UV crosslinking<\/strong><\/a>, in which adjacent thymine bases bind with one another (Figure 5.4). Many of these mutations are detected and corrected by <strong>DNA repair mechanisms<\/strong>, enzymes that patrol and repair DNA in living cells, while other mutations may cause a new freckle or mole or, perhaps, an unusual hair to grow. For people with the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1484\">autosomal recessive<\/a><\/strong> disease <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1486\"><strong>xeroderma pigmentosum<\/strong><\/a>, these repair mechanisms do not function correctly, resulting in a host of problems especially related to sun exposure, including severe sunburns, dry skin, heavy freckling, and other pigment changes.<\/p>\n<p class=\"import-Normal\">Most of our mutations exist in <strong>somatic<\/strong> cells, which are the cells of our organs and other body tissues. Those will not be passed onto future generations and so will not affect the population over time. Only mutations that occur in the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_686\">gametes<\/a><\/strong>, the reproductive cells (i.e., the sperm or egg cells), will be passed onto future generations. When a new mutation pops up at random in a family lineage, it is known as a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1490\"><strong>spontaneous mutation<\/strong><\/a>. If the individual born with this spontaneous mutation passes it on to his offspring, those offspring receive an <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1492\">inherited mutation<\/a><\/strong>. Geneticists have identified many classes of mutations and the causes and effects of many of these.<\/p>\n<h4 class=\"import-Normal\"><em>Point Mutations<\/em><\/h4>\n<p class=\"import-Normal\">A <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1494\">point mutation<\/a><\/strong> is a single-letter (single-nucleotide) change in the genetic code resulting in the substitution of one nucleic acid base for a different one. As you learned in Chapter 4, the DNA code in each gene is translated through three-letter \u201cwords\u201d known as <strong>codons<\/strong>. So depending on how the point mutation changes the \u201cword,\u201d the effect it will have on the protein may be major or minor or may make no difference at all.<\/p>\n<p class=\"import-Normal\">If a mutation does not change the resulting protein, then it is called a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1496\">synonymous mutation<\/a><\/strong>. Synonymous mutations do involve a letter (nucleic acid) change, but that change results in a codon that codes for the same \u201cinstruction\u201d (the same amino acid or stop code) as the original codon. Mutations that do cause a change in the protein are known as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1498\"><strong>nonsynonymous mutations<\/strong>.<\/a> Nonsynonymous mutations may change the resulting protein\u2019s amino acid sequence by altering the DNA sequence that encodes the mRNA or by changing how the mRNA is spliced prior to translation (refer to Chapter 4 for more details).<\/p>\n<h4 class=\"import-Normal\"><em>Insertions and Deletions<\/em><\/h4>\n<p class=\"import-Normal\">In addition to point mutations, another class of mutations are <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1500\"><strong>insertions<\/strong><\/a> and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1504\">deletions<\/a><\/strong>, or <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1502\"><strong>indels<\/strong><\/a>, for short. As the name suggests, these involve the addition (insertion) or removal (deletion) of one or more coding sequence letters (nucleic acids). These typically first occur as an error in DNA replication, wherein one or more nucleotides are either duplicated or skipped in error. Entire codons or sets of codons may also be removed or added if the indel is a multiple of three nucleotides.<\/p>\n<p class=\"import-Normal\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1506\"><strong>Frameshift<\/strong> <strong>mutations<\/strong><\/a> are types of indels that involve the insertion or deletion of any number of nucleotides that is not a multiple of three (e.g., adding one or two extra letters to the code). Because these indels are not consistent with the codon numbering, they \u201cshift the reading frame,\u201d causing all the codons beyond the mutation to be misread. Like point mutations, small indels can also disrupt splice sites.<\/p>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1508\">Transposable elements<\/a><\/strong>, or <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1510\">transposons<\/a><\/strong>, are fragments of DNA that can \u201cjump\u201d around in the genome. There are two types of transposons: <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1512\">retrotransposons<\/a><\/strong> are transcribed from DNA into RNA and then \u201creverse transcribed,\u201d to insert the copied sequence into a new location in the DNA, and<strong> <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1514\">DNA transposons<\/a><\/strong>, which do not involve RNA. DNA transposons are clipped out of the DNA sequence itself and inserted elsewhere in the genome. Because transposable elements insert themselves into existing DNA sequences, they are frequent gene disruptors. At certain times, and in certain species, it appears that transposons became very active, likely accelerating the mutation rate (and thus, the genetic variation) in those populations during the active periods.<\/p>\n<h4 class=\"import-Normal\"><em>Chromosomal Alterations <\/em><\/h4>\n<p class=\"import-Normal\">The final major category of genetic mutations are changes at the chromosome level: crossover events, nondisjunction events, and translocations. <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1516\"><strong>Crossover events<\/strong> <\/a> occur when DNA is swapped between homologous chromosomes while they are paired up during meiosis I. Crossovers are thought to be so common that some DNA swapping may happen every time chromosomes go through meiosis I. Crossovers don\u2019t necessarily introduce new alleles into a population, but they do make it possible for new combinations of alleles to exist on a single chromosome that can be passed to future generations. This also enables new combinations of alleles to be found within siblings who share the same parents. Also, if the fragments that cross over don\u2019t break at exactly the same point, they can cause genes to be deleted from one of the homologous chromosomes and duplicated on the other.<\/p>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1518\">Nondisjunction events<\/a><\/strong> occur when the homologous chromosomes (in meiosis I) or sister chromatids (in meiosis II and mitosis) fail to separate after pairing. The result is that both chromosomes or chromatids end up in the same daughter cell, leaving the other daughter cell without any copy of that chromosome (Figure 5.5). Most nondisjunctions at the gamete level are fatal to the embryo. The most widely known exception is Trisomy 21, or Down syndrome, which results when an embryo inherits three copies of Chromosome 21: two from one parent (due to a nondisjunction event) and one from the other (Figure 5.6). <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1520\">Trisomies<\/a> <\/strong>(triple chromosome conditions) of Chromosomes 18 (Edwards syndrome) and 13 (Patau syndrome) are also known to result in live births, but the children usually have severe complications and rarely survive beyond the first year of life.<\/p>\n<figure style=\"width: 601px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5.jpg\" alt=\"Egg cell undergoes normal meiosis and nondisjunction in meisosis 1.\" width=\"601\" height=\"391\" \/><figcaption class=\"wp-caption-text\">Figure 5.5: Illustration of an egg cell (oocyte) undergoing normal meiosis 1, resulting in a diploid daughter cell, compared to an egg cell undergoing nondisjunction during meiosis 1, resulting in a trisomy in the daughter cell. Credit: <a class=\"rId35\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Trisomy_due_to_nondisjunction_in_maternal_meiosis_1.png\">Trisomy due to nondisjunction in maternal meiosis 1<\/a> by Wpeissner has been modified (labels deleted by Katie Nelson) and is under a <a class=\"rId36\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<figure style=\"width: 316px\" class=\"wp-caption alignright\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image6-1.jpg\" alt=\"A young woman in a blue polo shirt smiles at the camera.\" width=\"316\" height=\"364\" \/><figcaption class=\"wp-caption-text\">Figure 5.6: Amy Bockerstette, a competitive golfer and disabilities advocate, also has Down Syndrome. Credit: <a class=\"rId38\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Amy_Bockerstette_Headshot.jpg\">Amy Bockerstette Headshot<\/a> by Bucksgrandson is under a <a class=\"rId39\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Sex chromosome trisomies (XXX, XXY, XYY) and X chromosome <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1522\"><strong>monosomies<\/strong> <\/a>(inheritance of an X chromosome from one parent and no sex chromosome from the other) are also survivable and fairly common. The symptoms vary but often include atypical sexual characteristics, either at birth or at puberty, and often result in sterility. The X chromosome carries unique genes that are required for survival; therefore, Y chromosome monosomies are incompatible with life.<\/p>\n<p class=\"import-Normal\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1524\"><strong>Chromosomal translocations<\/strong><\/a> involve transfers of DNA between nonhomologous chromosomes. This may involve swapping large portions of two or more chromosomes. The exchanges of DNA may be balanced or unbalanced. In <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1526\"><strong>balanced translocations<\/strong><\/a>, the genes are swapped, but no genetic information is lost. In <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1528\">unbalanced translocations<\/a><\/strong>, there is an unequal exchange of genetic material, resulting in duplication or loss of genes. Translocations result in new chromosomal structures called <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1530\"><strong>derivative chromosomes<\/strong><\/a>, because they are derived or created from two different chromosomes<em>. <\/em>Translocations are often found to be linked to cancers and can also cause infertility. Even if the translocations are balanced in the parent, the embryo often won\u2019t survive unless the baby inherits both of that parent\u2019s derivative chromosomes (to maintain the balance).<\/p>\n<h3 class=\"import-Normal\"><strong>Genetic Drift<\/strong><\/h3>\n<p class=\"import-Normal\">The second force of evolution is commonly known as genetic drift. This is an unfortunate misnomer, as this force actually involves the drifting of alleles, not genes. <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1532\"><strong>Genetic <\/strong><strong>d<\/strong><strong>rift<\/strong><\/a> refers to <em>random<\/em> changes (\u201cdrift\u201d) in allele frequencies from one generation to the next. The genes are remaining constant within the population; it is only the alleles of the genes that are changing in frequency. The random nature of genetic drift is a crucial point to understand: it specifically occurs when none of the variant alleles confer an advantage.<\/p>\n<figure style=\"width: 368px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image7-2.png\" alt=\"A smooth cell has a gently curving exterior surface, and a ruffled cell has undulating surface.\" width=\"368\" height=\"215\" \/><figcaption class=\"wp-caption-text\">Figure 5.7: Smooth and ruffled amoeba-like cells. Credit: Smooth and ruffled amoeba-like cells original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Andrea J. Alveshere is a collective work under a <a class=\"rId41\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Includes <a class=\"rId42\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Amoeba_proteus_TK-UT.svg\">Amoeba Proteus TK-UT<\/a> by <a class=\"rId43\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Nefronus\">Tom\u00e1\u0161 Kebert<\/a> and <a class=\"rId44\" href=\"https:\/\/www.umimeto.org\/\">umimeto.org<\/a> (modified), <a class=\"rId45\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0<\/a>.]<\/figcaption><\/figure>\n<p class=\"import-Normal\">Let\u2019s imagine far back in time, again, to that ancient population of amoeba-like cells, subsisting and occasionally dividing, in the primordial sea. A mutation occurs in one of the cells that changes the texture of the cell membrane from a relatively smooth surface to a highly ruffled one (Figure 5.7). This has absolutely no effect on the cell\u2019s quality of life or ability to reproduce. In fact, eyes haven\u2019t evolved yet, so no one in the world at the time would even notice the difference. The cells in the population continue to divide, and the offspring of the ruffled cell inherit the ruffled membrane. The frequency (percentage) of the ruffled allele in the population, from one generation to the next, will depend entirely on how many offspring that first ruffled cell ends up having, and the random events that might make the ruffled alleles more common or more rare (such as population bottlenecks and founder effects, which are discussed below).<\/p>\n<h4 class=\"import-Normal\"><em>Sexual Reproduction and Random Inheritance<\/em><\/h4>\n<p class=\"import-Normal\">Tracking alleles gets a bit more complicated in our primordial cells when, after a number of generations, a series of mutations have created populations that reproduce sexually. These cells now must go through an extra round of cell division (meiosis) to create haploid gametes. The combination of two gametes is now required to produce each new diploid offspring.<\/p>\n<figure style=\"width: 262px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image8-1.png\" alt=\"A Punnett square with ruffled and smooth cells.\" width=\"262\" height=\"262\" \/><figcaption class=\"wp-caption-text\">Figure 5.8: A Punnett square demonstrating the sexual inheritance pattern of ruffled (dominant) and smooth amoeba-like primordial cells. Credit: Punnett square of primordial cells original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Andrea J. Alveshere is a collective work under a <a class=\"rId47\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Includes <a class=\"rId48\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Amoeba_proteus_TK-UT.svg\">Amoeba Proteus TK-UT<\/a> by <a class=\"rId49\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Nefronus\">Tom\u00e1\u0161 Kebert<\/a> and <a class=\"rId50\" href=\"https:\/\/www.umimeto.org\/\">umimeto.org<\/a> (modified), <a class=\"rId51\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0<\/a>; <a class=\"rId52\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Punnett_hetero_x_hetero.svg\">Punnett Hetero x Hetero<\/a> by <a class=\"rId53\" href=\"https:\/\/commons.wikimedia.org\/w\/index.php?title=User:Purpy_Pupple&amp;redirect=no\">Purpy Pupple<\/a> (modified), <a class=\"rId54\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/deed.en\">CC BY-SA 3.0<\/a>].<\/figcaption><\/figure>\n<p class=\"import-Normal\">In the earlier population, which reproduced via <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1534\">asexual reproduction<\/a><\/strong>, a cell either carried the smooth allele or the ruffled allele. With <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1536\">sexual reproduction<\/a><\/strong>, a cell inherits one allele from each parent, so there are homozygous cells that contain two smooth alleles, homozygous cells that contain two ruffled alleles, and heterozygous cells that contain one of each allele (Figure 5.8). If the new, ruffled allele happens to be dominant (and we\u2019ll imagine that it is), the heterozygotes will have ruffled cell <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1238\"><strong>phenotypes<\/strong><\/a> but also will have a 50\/50 chance of passing on a smooth allele to each offspring. As long as neither phenotype (ruffled nor smooth) provides any advantage over the other, the variation in the population from one generation to the next will remain completely random.<\/p>\n<p class=\"import-Normal\">In sexually reproducing populations (including humans and many other animals and plants in the world today), that 50\/50 chance of inheriting one or the other allele from each parent plays a major role in the random nature of genetic drift.<\/p>\n<h4 class=\"import-Normal\"><em>Population Bottlenecks <\/em><\/h4>\n<p class=\"import-Normal\">A <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1538\"><strong>population bottleneck<\/strong><\/a> occurs when the number of individuals in a population drops dramatically due to some random event. The most obvious, familiar examples are natural disasters. Tsunamis and hurricanes devastating island and coastal populations and forest fires and river floods wiping out populations in other areas are all too familiar. When a large portion of a population is randomly wiped out, the allele frequencies (i.e., the percentages of each allele) in the small population of survivors are often much different from the frequencies in the predisaster, or \u201cparent,\u201d population.<\/p>\n<p class=\"import-Normal\">If such an event happened to our primordial ocean cell population\u2014perhaps a volcanic fissure erupted in the ocean floor and only the cells that happened to be farthest from the spewing lava and boiling water survived\u2014we might end up, by random chance, with a surviving population that had mostly ruffled alleles, in contrast to the parent population, which had only a small percentage of ruffles (Figure 5.9).<\/p>\n<figure style=\"width: 665px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image9-2.png\" alt=\"Ruffled and smooth cells experience population bottleneck when a lava flow divides the populations.\" width=\"665\" height=\"332\" \/><figcaption class=\"wp-caption-text\">Figure 5.9: Illustration of a population of amoeba-like cells shifting from primarily smooth phenotypes (at left) to mostly ruffled phenotypes due to eruption of a volcanic fissure (at right) that exterminated the nearest cells. Credit: Population of amoeba-like cells and volcanic fissure original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Andrea J. Alveshere is a collective work under a <a class=\"rId56\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Includes <a class=\"rId57\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Amoeba_proteus_TK-UT.svg\">Amoeba Proteus TK-UT<\/a> by <a class=\"rId58\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Nefronus\">Tom\u00e1\u0161 Kebert<\/a> and <a class=\"rId59\" href=\"https:\/\/www.umimeto.org\/\">umimeto.org<\/a> (modified), <a class=\"rId60\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0<\/a>.]<\/figcaption><\/figure>\n<p class=\"import-Normal\">One of the most famous examples of a population bottleneck is the prehistoric disaster that led to the extinction of dinosaurs, the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1540\"><strong>Cretaceous\u2013Paleogene <\/strong><strong>extinction<\/strong><\/a> event (often abbreviated K\u2013Pg; previously K-T). This occurred approximately 66 million years ago. Dinosaurs and all their neighbours were going about their ordinary routines when a massive asteroid zoomed in from space and crashed into what is now the Gulf of Mexico, creating an impact so enormous that populations within hundreds of miles of the crash site were likely immediately wiped out. The skies filled with dust and debris, causing temperatures to plummet worldwide. It\u2019s estimated that 75% of the world\u2019s species went extinct as a result of the impact and the deep freeze that followed (Jablonski and Chaloner 1994).<\/p>\n<figure style=\"width: 399px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image10-2.png\" alt=\" A rat-like creature sits atop a dinosaur, raising a fist in a victorious gesture.\" width=\"399\" height=\"323\" \/><figcaption class=\"wp-caption-text\">Figure 5.10: The Cretaceous\u2013Paleogene extinction event, which led to the fall of the dinosaurs and rise of the mammals. Credit: <a class=\"rId62\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">The<\/a> <a class=\"rId64\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Cretaceous\u2013Paleogene extinction event (Figure 4.12)<\/a> original to <a class=\"rId65\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is 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\">The populations that emerged from the K-Pg extinction were markedly different from their pre-disaster communities. Surviving mammal populations expanded and diversified, and other new creatures appeared. The ecosystems of Earth were filled with new organisms and have never been the same (Figure 5.10).<\/p>\n<p class=\"import-Normal\">Much more recently in geological time, during the colonial period, many human populations experienced bottlenecks as a result of the fact that imperial powers were inclined to slaughter communities who were reluctant to give up their lands and resources. This effect was especially profound in the Americas, where Indigenous populations faced the compounded effects of brutal warfare, exposure to new bacteria and viruses (against which they had no immunity), and ultimately segregation on resource-starved reservations. The populations in Europe, Asia, and Africa had experienced regular gene flow during the 10,000-year period in which most kinds of livestock were being domesticated, giving them many generations of experience building up immunity against zoonotic diseases (those that can pass from animals to humans). In contrast, the residents of the Americas had been almost completely isolated during those millennia, so all these diseases swept through the Americas in rapid succession, creating a major loss of genetic diversity in the Indigenous American population. It is estimated that between 50% and 95% of the Indigenous American populations died during the first decades after European contact, around 500 years ago (Livi-Bacci 2006).<\/p>\n<p class=\"import-Normal\">An urgent health challenge facing humans today involves human-induced population bottlenecks that produce antibiotic-resistant bacteria. <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1451\"><strong>Antibiotics<\/strong> <\/a>are medicines prescribed to treat bacterial infections. The typical prescription includes enough medicine for ten days. People often feel better much sooner than ten days and sometimes decide to quit taking the medicine ahead of schedule. This is often a big mistake. The antibiotics have quickly killed off a large percentage of the bacteria\u2014enough to reduce the symptoms and make you feel much better. However, this has created a bacterial population bottleneck. There are usually a small number of bacteria that survive those early days. If you take the medicine as prescribed for the full ten days, it\u2019s quite likely that there will be no bacterial survivors. If you quit early, though, the survivors\u2014who were the members of the original population who were most resistant to the antibiotic\u2014will begin to reproduce again. Soon the infection will be back, possibly worse than before, and now all of the bacteria are resistant to the antibiotic that you had been prescribed.<\/p>\n<p class=\"import-Normal\">Other activities that have contributed to the rise of antibiotic-resistant bacteria include the use of antibacterial cleaning products and the inappropriate use of antibiotics as a preventative measure in livestock or to treat infections that are viral instead of bacterial (viruses do not respond to antibiotics). In 2017, the World Health Organization published a list of twelve antibiotic-resistant pathogens that are considered top priority targets for the development of new antibiotics (World Health Organization 2017).<\/p>\n<div class=\"textbox shaded\" style=\"background: var(--lightblue)\">\n<h2>Dig Deeper: The North American Elephant Seal: Thriving Bottleneck Populations That Still Face Genetic Defects<\/h2>\n<p>In 1892, the Northern Elephant Seal underwent a severe population bottleneck caused by commercial hunting, reducing the species to an estimated 20 individuals at the time. This drastic decline led to a substantial loss of genetic diversity\u2013a common consequence of extreme population bottlenecks (Hoelzel Et al., 2024 &amp; Weber Et al., 2000). While the population has since recovered to over 200,000 individuals, its genetic variability remains significantly low. Analyses of genetic markers, including allozymes, mitochondrial DNA, and microsatellites, consistently reflect this reduced diversity (Hoelzel Et al., 2024). Comparative studies further underscore this loss by highlighting the higher genetic variation observed in the Southern Elephant Seal, which did not experience similar population constraints (2024).<\/p>\n<figure style=\"width: 386px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/upload.wikimedia.org\/wikipedia\/commons\/thumb\/4\/48\/Elephant_seals_at_Ano_Nuevo_%2891577%29.jpg\/250px-Elephant_seals_at_Ano_Nuevo_%2891577%29.jpg\" alt=\"File:Elephant seals at Ano Nuevo (91577).jpg\" width=\"386\" height=\"295\" \/><figcaption class=\"wp-caption-text\">Figure 5.11 A male northern elephant seal (Mirounga angustirostris) with two pups at Ano Nuevo State Park. Credit: Elephant seals at Ano Nuevo by Rhododendrites is under <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\" target=\"_blank\" rel=\"noopener\">Creative Commons Attribution-Share Alike 4.0<\/a>.<\/figcaption><\/figure>\n<p>In a 2024 study for Nature, Ecology, and Evolution, Hoelzel and colleagues sequenced 260 modern and 8 historical genomes of the northern elephant seal. This comparison revealed a decrease in average heterozygosity from 0.00142 before the bottleneck to 0.000176 in the contemporary population, confirming the decline in genetic variation (2024). Hoelzel\u2019s mitogenome tree further illustrates this loss, revealing only two significant lineages remaining post-bottleneck, with limited diversity within each. Among the issues of diversity, the population has shown an increased number of loss-of-function (LOF) alleles, suggesting that increased inbreeding has amplified the frequency of these detrimental alleles; this reduced genetic diversity negatively affects both male and female reproductive fitness. Females who practiced repetitive inbreeding had higher LOF alleles and subsequently weaned fewer pups per year over their lifetime, while male reproductive success was linked to specific LOF loci associated with sperm production (2024). Hoelzel uses the example of \u201cAlpha-Male M12\u201d\u2013known for low paternity success despite frequent copulations\u2013which was homozygous for non-functional versions of four out of five LOF loci related to sperm function (2024, p. 688). The species' mating system, characterized by extreme polygyny, further exacerbates the loss of genetic variation even with countless copulatory partners<\/p>\n<p>Prior research published in Current Biology presents an empirical genetic assessment of this population bottleneck, highlighting its long-term genetic consequences, particularly the loss of mitochondrial diversity (Weber et al., 2000). In this research, Weber and colleagues note that random lineage sampling during the bottleneck led to the persistence of specific genetic variants by chance rather than through natural selection (2000). This research emphasizes that the loss of diversity poses potential future genetic vulnerabilities for the seals, and that further studies are crucial for understanding the full scope of these impacts on the seals' overall fitness (2000). In 2024, the work led by Hoelzen and company provided the missing data that the previous study had left unanswered. Their previously explored findings indicate that, although the seals have recovered in numbers, their genetic resilience remains compromised, leaving the population more vulnerable to future environmental pressures, such as climate change or resource shortages (Hoelzel Et al., 2024). Ultimately, while the population's size remains stable, the genetic consequences of the bottleneck indicate that past stochastic events continue to influence the seals' long-term fitness and adaptability.<\/p>\n<p>This research indicates that the historical bottleneck continues to affect the seals' health and fitness, despite the population's recovery. Limited genetic diversity and the persistence of harmful alleles due to inbreeding have continued to handicap the species' ability to thrive in environmental challenges such as climate change and resource fluctuations (2024). This emphasizes the importance of incorporating genetic factors into conservation strategies, as populations that have rebounded may still harbour long-term genetic weaknesses. Moreover, the elephant seal\u2019s history serves as a powerful example of how human actions \u2014such as overhunting \u2014 can have long-lasting impacts on biodiversity, reinforcing the importance of understanding human-environment interactions in ecological and conservation contexts.<\/p>\n<\/div>\n<h4 class=\"import-Normal\"><em>Founder Effects<\/em><\/h4>\n<p class=\"import-Normal\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1452\"><strong>Founder effects<\/strong><\/a> occur when members of a population leave the main or \u201cparent\u201d group and form a new population that no longer interbreeds with the other members of the original group. Similar to survivors of a population bottleneck, the newly founded population often has allele frequencies that are different from the original group. Alleles that may have been relatively rare in the parent population can end up being very common due to the founder effect. Likewise, recessive traits that were seldom seen in the parent population may be seen frequently in the descendants of the offshoot population.<\/p>\n<p class=\"import-Normal\">One striking example of the founder effect was first noted in the Dominican Republic in the 1970s. During a several-year period, eighteen children who had been born with female genitalia and raised as girls suddenly grew penises at puberty. This culture tended to value sons over daughters, so these transitions were generally celebrated. They labeled the condition <em><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1453\">guevedoces<\/a><\/strong><\/em>, which translates to \u201cpenis at twelve,\u201d due to the average age at which this occurred. Scientists were fascinated by the phenomenon.<\/p>\n<p class=\"import-Normal\">Genetic and hormonal studies revealed that the condition, scientifically termed <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1454\">5-alpha reductase deficiency<\/a>,<\/strong> is an autosomal recessive syndrome that manifests when a child having both X and Y sex chromosomes inherits two nonfunctional (mutated) copies of the <em>SRD5A2 <\/em>gene (Imperato-McGinley &amp; Zhu 2002). These children develop testes internally, but the 5-alpha reductase 2 steroid, which is necessary for development of male genitals in babies, is not produced. In absence of this male hormone, the baby develops female-looking genitalia (in humans, \u201cfemale\u201d is the default infant body form, if the full set of the necessary male hormones are not produced). At puberty, however, a different set of male hormones are produced by other fully functional genes. These hormones complete the male genital development that did not happen in infancy. This condition became quite common in the Dominican Republic during the 1970s due to founder effect\u2014that is, the mutated <em>SRD5A2<\/em> gene happened to be much more common among the Dominican Republic\u2019s founding population than in the parent populations. (The Dominican population derives from a mixture of Indigenous Americans [Taino] peoples, West Africans, and Western Europeans.) Five-alpha reductase syndrome has since been observed in other small, isolated populations around the world.<\/p>\n<p class=\"import-Normal\">Founder effect is closely linked to the concept of inbreeding, which in population genetics does not necessarily mean breeding with immediate family relatives. Instead, <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1455\"><strong>inbreeding<\/strong> <\/a> refers to the selection of mates exclusively from within a small, closed population\u2014that is, from a group with limited allelic variability. This can be observed in small, physically isolated populations but also can happen when cultural practices limit mates to a small group. As with the founder effect, inbreeding increases the risk of inheriting two copies of any nonfunctional (mutant) alleles.<\/p>\n<p class=\"import-Normal\">The Amish in the United States are a population that, due to their unique history and cultural practices, emerged from a small founding population and have tended to select mates from within their groups. The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1456\">Old Order Amish<\/a><\/strong> population of Lancaster County, Pennsylvania, has approximately 50,000 current members, all of whom can trace their ancestry back to a group of approximately 80 individuals. This small founding population immigrated to the United States from Switzerland in the mid-1700s to escape religious persecution. Since the Amish keep to themselves and almost exclusively select mates from within their own communities, they have more recessive traits compared to their parent population.<\/p>\n<figure style=\"width: 441px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image11-1.jpg\" alt=\"One individual\u2019s hands with six fingers.\" width=\"441\" height=\"331\" \/><figcaption class=\"wp-caption-text\">Figure 5.12: A person displaying polydactyly. Credit: <a class=\"rId68\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:6_Finger.JPG\">6 Finger<\/a> by Wilhelmy is under a <a class=\"rId69\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">One of the genetic conditions that has been observed much more frequently in the Lancaster County Amish population is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1457\">Ellis-van Creveld syndrome<\/a><\/strong>, which is an autosomal recessive disorder characterized by short stature (dwarfism), polydactyly (the development of more than five digits [fingers or toes] on the hands or feet], abnormal tooth development, and heart defects (Figure 5.12). Among the general world population, Ellis-van Creveld syndrome is estimated to affect approximately 1 in 60,000 individuals; among the Old Order Amish of Lancaster County, the rate is estimated to be as high as 1 in every 200 births (D\u2019Asdia Et al. 2013).<\/p>\n<p class=\"import-Normal\">One important insight that has come from the study of founder effects is that a limited gene pool carries a much higher risk for genetic diseases. Genetic diversity in a population greatly reduces these risks.<\/p>\n<h3 class=\"import-Normal\"><strong>Gene Flow<\/strong><\/h3>\n<p class=\"import-Normal\">The third force of evolution is traditionally called gene flow. As with genetic drift, this is a misnomer, because it refers to flowing alleles, not genes. (All members of the same species share the same genes; it is the alleles of those genes that may vary.) <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1459\"><strong>Gene <\/strong><strong>f<\/strong><strong>low<\/strong> <\/a> refers to the movement of alleles from one population to another. In most cases, gene flow can be considered synonymous with migration.<\/p>\n<p class=\"import-Normal\">Returning again to the example of our primordial cell population, let\u2019s imagine that, after the volcanic fissure opened up in the ocean floor, wiping out the majority of the parent population, two surviving populations developed in the waters on opposite sides of the fissure. Ultimately, the lava from the fissure cooled into a large island that continued to provide a physical barrier between the populations (Figure 5.13).<\/p>\n<figure style=\"width: 685px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image12-2.png\" alt=\"An illustration of gene flow.\" width=\"685\" height=\"342\" \/><figcaption class=\"wp-caption-text\">Figure 5.13: Smooth and predominantly ruffled amoeba-like populations separated by a volcanic eruption (at left) and an island (at right) with unidirectional gene flow moving from east to west with ocean currents. Credit: Population of amoeba-like cells separated by volcanic eruption original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Andrea J. Alveshere is a collective work under a <a class=\"rId74\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Includes <a class=\"rId75\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Amoeba_proteus_TK-UT.svg\">Amoeba Proteus TK-UT<\/a> by <a class=\"rId76\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Nefronus\">Tom\u00e1\u0161 Kebert<\/a> and <a class=\"rId77\" href=\"https:\/\/www.umimeto.org\/\">umimeto.org<\/a> (modified), <a class=\"rId78\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0<\/a>.]<\/figcaption><\/figure>\n<p class=\"import-Normal\">In the initial generations after the eruption, due to founder effect, isolation, and random inheritance (genetic drift), the population to the west of the islands contained a vast majority of the ruffled membrane alleles while the eastern population carried only the smooth alleles. Ocean currents in the area typically flowed from east to west, sometimes carrying cells (facilitating gene flow) from the eastern (smooth) population to the western (ruffled) population. Due to the ocean currents, it was almost impossible for any cells from the western population to be carried eastward. Thus, for inheritance purposes, the eastern (smooth) population remained isolated. In this case, the gene flow is unidirectional (going only in one direction) and unbalanced (only one population is receiving the new alleles).<\/p>\n<p class=\"import-Normal\">Among humans, gene flow is often described as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1460\"><strong>admixture<\/strong>.<\/a> In forensic cases, anthropologists and geneticists are often asked to estimate the ancestry of unidentified human remains to help determine whether they match any missing persons\u2019 reports. This is one of the most complicated tasks in these professions because, while \u201crace\u201d or \u201cancestry\u201d involves simple checkboxes on a missing person\u2019s form, among humans today there are no truly distinct genetic populations. All modern humans are members of the same fully breeding compatible species, and all human communities have experienced multiple episodes of gene flow (admixture), leading all humans today to be so genetically similar that we are all members of the same (and only surviving) human subspecies: <em>Homo sapiens sapiens.<\/em><\/p>\n<p class=\"import-Normal\">Gene flow between otherwise isolated nonhuman populations is often termed <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1461\">hybridization.<\/a>.<\/strong> One example of this involves the hybridization and spread of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1462\"><strong>Scutellata<\/strong><strong> honey bees<\/strong><\/a> (a.k.a. \u201ckiller bees\u201d) in the Americas. All honey bees worldwide are classified as <em>Apis mellifera.<\/em> Due to distinct adaptations to various environments around the world, there are 28 different subspecies of <em>Apis mellifera<\/em>.<\/p>\n<p class=\"import-Normal\">During the 1950s, a Brazilian biologist named Warwick E. Kerr experimented with hybridizing African and European subspecies of honey bees to try to develop a strain that was better suited to tropical environments than the European honey bees that had long been kept by North American beekeepers. Dr. Kerr was careful to contain the reproductive queens and drones from the African subspecies, but in 1957, a visiting beekeeper accidentally released 26 queen bees of the Scutellata subspecies (<em>Apis mellifera scutellata<\/em>) from southern Africa into the Brazilian countryside. The Scutellata bees quickly interbred with local European honey bee populations. The hybridized bees exhibited a much more aggressively defensive behaviour, fatally or near-fatally attacking many humans and livestock that ventured too close to their hives. The hybridized bees spread throughout South America and reached Mexico and California by 1985. By 1990, permanent colonies had been established in Texas, and by 1997, 90% of trapped bee swarms around Tucson, Arizona, were found to be Scutellata hybrids (Sanford 2006).<\/p>\n<p class=\"import-Normal\">Another example involves the introduction of the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1463\"><strong>Harlequin ladybeetle<\/strong><\/a>, <em>Harmonia axyridis<\/em>, native to East Asia, to other parts of the world as a \u201cnatural\u201d form of pest control. Harlequin ladybeetles are natural predators of some of the aphids and other crop-pest insects. First introduced to North America in 1916, the \u201cbiocontrol\u201d strains of Harlequin ladybeetles were considered to be quite successful in reducing crop pests and saving farmers substantial amounts of money. After many decades of successful use in North America, biocontrol strains of Harlequin ladybeetles were also developed in Europe and South America in the 1980s.<\/p>\n<p class=\"import-Normal\">Over the seven decades of biocontrol use, the Harlequin ladybeetle had never shown any potential for development of wild colonies outside of its native habitat in China and Japan. New generations of beetles always had to be reared in the lab. That all changed in 1988, when a wild colony took root near New Orleans, Louisiana. Either through admixture with a native ladybeetle strain, or due to a spontaneous mutation, a new allele was clearly introduced into this population that suddenly enabled them to survive and reproduce in a wide range of environments. This population spread rapidly across the Americas and had reached Africa by 2004.<\/p>\n<p class=\"import-Normal\">In Europe, the invasive, North American strain of Harlequin ladybeetle admixed with the European strain (Figure 5.14), causing a population explosion (Lombaert Et al. 2010). Even strains specifically developed to be flightless (to curtail the spreading) produced flighted offspring after admixture with members of the North American population (Facon Et al. 2011). The fast-spreading, invasive strain has quickly become a disaster, out-competing native ladybeetle populations (some to the point of extinction), causing home infestations, decimating fruit crops, and contaminating many batches of wine with their bitter flavour after being inadvertently harvested with the grapes (Pickering Et al. 2004).<\/p>\n<figure style=\"width: 583px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image13-2.png\" alt=\"One gray ladybug is migrating to the group of white ladybugs.\" width=\"583\" height=\"219\" \/><figcaption class=\"wp-caption-text\">Figure 5.14: Gene flow between two populations of ladybeetles (ladybugs). Credit: <a class=\"rId80\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Ladybug Gene Flow (Figure 4.14)<\/a> original to <a class=\"rId81\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a class=\"rId82\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Natural Selection<\/strong><\/h3>\n<p class=\"import-Normal\">The final force of evolution is natural selection. This is the evolutionary process that Charles Darwin first brought to light, and it is what the general public typically evokes when considering the process of evolution. <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1465\"><strong>Natural <\/strong><strong>s<\/strong><strong>election<\/strong><\/a> occurs when certain phenotypes confer an advantage or disadvantage in survival and\/or reproductive success. The alleles associated with those phenotypes will change in frequency over time due to this selective pressure. It\u2019s also important to note that the advantageous allele may change over time (with environmental changes) and that an allele that had previously been benign may become advantageous or detrimental. Of course, dominant, recessive, and codominant traits will be selected upon a bit differently from one another. Because natural selection acts on phenotypes rather than the alleles themselves, deleterious (disadvantageous) alleles can be retained by heterozygotes without any negative effects.<\/p>\n<p class=\"import-Normal\">In the case of our primordial ocean cells, up until now, the texture of their cell membranes has been benign. The frequencies of smooth to ruffled alleles, and smooth to ruffled phenotypes, has changed over time, due to genetic drift and gene flow. Let\u2019s now imagine that the Earth\u2019s climate has cooled to a point that the waters frequently become too cold for survival of the tiny bacteria that are the dietary staples of our smooth and ruffled cell populations. The way amoeba-like cells \u201ceat\u201d is to stretch out the cell membrane, almost like an arm, to encapsulate, then ingest, the tiny bacteria. When the temperatures plummet, the tiny bacteria populations plummet with them. Larger bacteria, however, are better able to withstand the temperature change.<\/p>\n<p class=\"import-Normal\">The smooth cells were well-adapted to ingesting tiny bacteria but poorly suited to encapsulating the larger bacteria. The cells with the ruffled membranes, however, are easily able to extend their ruffles to encapsulate the larger bacteria. They also find themselves able to stretch their entire membrane to a much larger size than their smooth-surfaced neighbours, allowing them to ingest more bacteria at a given time and to go for longer periods between feedings (Figure 5.15).<\/p>\n<figure style=\"width: 528px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image14-2.png\" alt=\"Smooth and ruffled cells feeding on large and small bacteria.\" width=\"528\" height=\"307\" \/><figcaption class=\"wp-caption-text\">Figure 5.15: Smooth and ruffled cells feeding. Credit: Smooth and ruffled cells feeding original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Andrea J. Alveshere is a collective work under a <a class=\"rId84\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Includes <a class=\"rId85\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Cladograma_dos_Dom%C3%ADnios_e_Reinos.png\">Cladograma dos Dominios e Reinos<\/a> by <a class=\"rId86\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:MarceloTeles\">MarceloTeles<\/a> (modified), <a class=\"rId87\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0<\/a><a class=\"rId88\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">; <\/a><a class=\"rId89\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Amoeba_proteus_TK-UT.svg\">Amoeba Proteus TK-UT<\/a> by <a class=\"rId90\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Nefronus\">Tom\u00e1\u0161 Kebert<\/a> and <a class=\"rId91\" href=\"https:\/\/www.umimeto.org\/\">umimeto.org<\/a> (modified), <a class=\"rId92\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0<\/a>.]<\/figcaption><\/figure>\n<p class=\"import-Normal\">The smooth and ruffled traits, which had previously offered no advantage or disadvantage while food was plentiful, now are subject to natural selection. During the cold snaps, at least, the ruffled cells have a definite advantage. We can imagine that the western population that has mostly ruffled alleles will continue to do well, while the eastern population is at risk of dying out if the smaller bacteria remain scarce and no ruffled alleles are introduced.<\/p>\n<p class=\"import-Normal\">A classic example of natural selection involves the study of an insect called the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1466\">peppered moth<\/a><\/strong> (<em>Biston betularia<\/em>) in England during the Industrial Revolution in the 1800s. Prior to the Industrial Revolution, the peppered moth population was predominantly light in colour, with dark (pepper-like) speckles on the wings. The \u201cpeppered\u201d colouration was very similar to the appearance of the bark and lichens that grew on the local trees (Figure 5.16). This helped to camouflage the moths as they rested on a tree, making it harder for moth-eating birds to find and snack on them. There was another phenotype that popped up occasionally in the population. These individuals were heterozygotes that carried an overactive, dominant pigment allele, producing a solid black colouration. As you can imagine, the black moths were much easier for birds to spot, making this phenotype a real disadvantage.<\/p>\n<p class=\"import-Normal\">The situation changed, however, as the Industrial Revolution took off. Large factories began spewing vast amounts of coal smoke into the air, blanketing the countryside, including the lichens and trees, in black soot. Suddenly, it was the light-coloured moths that were easy for birds to spot and the black moths that held the advantage. The frequency of the dark pigment allele rose dramatically. By 1895, the black moth phenotype accounted for 98% of observed moths (Grant 1999).<\/p>\n<figure style=\"width: 476px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image15-2.png\" alt=\"An illustration of natural selection.\" width=\"476\" height=\"531\" \/><figcaption class=\"wp-caption-text\">Figure 5.16: Dark and light peppered moth variants and their relative camouflage abilities on clean (top) and sooty (bottom) trees. Credit: <a class=\"rId94\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Peppered_moths_c2.jpg\">Peppered moths c2<\/a> by Khaydock is under a <a class=\"rId95\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Thanks to new environmental regulations in the 1960s, the air pollution in England began to taper off. As the soot levels decreased, returning the trees to their former, lighter colour, this provided the perfect opportunity to study how the peppered moth population would respond. Repeated follow-up studies documented the gradual rise in the frequency of the lighter-coloured phenotype. By 2003, the maximum frequency of the dark phenotype was 50% and in most parts of England had decreased to less than 10% (Cook, 2003).<\/p>\n<h4 class=\"import-Normal\"><em>Directional, Balancing\/Stabilizing, and Disruptive\/Diversifying Selection<\/em><\/h4>\n<p class=\"import-Normal\">Natural selection can be classified as directional, balancing\/stabilizing, or disruptive\/diversifying, depending on how the pressure is applied to the population (Figure 5.17).<\/p>\n<figure style=\"width: 465px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image16-2.png\" alt=\"Three types of selection; balancing, directional and disruptive\/diversifying\" width=\"465\" height=\"574\" \/><figcaption class=\"wp-caption-text\">Figure 5.17: Lines depict the affects of (a) Balancing\/Stabilizing, (b) Directional, and (c) Disruptive\/Diversifying selection on populations. <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 class=\"rId97\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Figure_19_03_01.png\">Biology (ID: 185cbf87-c72e-48f5-b51e-f14f21b5eabd@9.17)<\/a> by <a class=\"rId98\" href=\"https:\/\/cnx.org\/\">CNX OpenStax<\/a> is used under a <a class=\"rId99\" href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/deed.en\">CC BY 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Both of the above examples of natural selection involve <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1467\">directional selection<\/a><\/strong>: the environmental pressures favour one phenotype over the other and cause the frequencies of the associated advantageous alleles (ruffled membranes, dark pigment) to gradually increase. In the case of the peppered moths, the direction shifted three times: first, it was selecting for lighter pigment; then, with the increase in pollution, the pressure switched to selection for darker pigment; finally, with reduction of the pollution, the selection pressure shifted back again to favouring light-coloured moths.<\/p>\n<p class=\"import-Normal\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1468\"><strong>Balancing selection<\/strong><\/a> (a.k.a. stabilizing selection) occurs when selection works against the extremes of a trait and favours the intermediate phenotype. For example, humans maintain an average birth weight that balances the need for babies to be small enough not to cause complications during pregnancy and childbirth but big enough to maintain a safe body temperature after they are born. Another example of balancing selection is found in the genetic disorder called sickle cell anemia (see \u201cSpecial Topic: Sickle Cell Anemia\u201d).<\/p>\n<p class=\"import-Normal\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1469\"><strong>Disruptive selection<\/strong><\/a> (a.k.a. diversifying selection), the opposite of balancing selection, occurs when both extremes of a trait are advantageous. Since individuals with traits in the mid-range are selected against, disruptive selection can eventually lead to the population evolving into two separate species. Darwin believed that the many species of finches (small birds) found in the remote Galapagos Islands provided a clear example of disruptive selection leading to speciation. He observed that seed-eating finches either had large beaks, capable of eating very large seeds, or small beaks, capable of retrieving tiny seeds. The islands did not have many plants that produced medium-size seeds. Thus, birds with medium-size beaks would have trouble eating the very large seeds and would also have been inefficient at picking up the tiny seeds. Over time, Darwin surmised, this pressure against mid-size beaks may have led the population to divide into two separate species.<\/p>\n<h4 class=\"import-Normal\"><em>Sexual Selection<\/em><\/h4>\n<p class=\"import-Normal\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1470\"><strong>Sexual <\/strong><strong>s<\/strong><strong>election<\/strong><\/a> is an aspect of natural selection in which the selective pressure specifically affects reproductive success (the ability to successfully breed and raise offspring) rather than survival. Sexual selection favours traits that will attract a mate. Sometimes these sexually appealing traits even carry greater risks in terms of survival.<\/p>\n<figure style=\"width: 354px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image17-2.png\" alt=\"A fox chases a peacock fleeing; a peacock displays his feathers to a peahen.\" width=\"354\" height=\"413\" \/><figcaption class=\"wp-caption-text\">Figure 5.18: Showy peacock tail disadvantages (becoming easier prey) and advantages (impressing peahens). Credit: <a class=\"rId101\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Peacock tail advantage and disadvantages (Figure 4.18)<\/a> original to <a class=\"rId102\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a class=\"rId103\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.License.<\/figcaption><\/figure>\n<p class=\"import-Normal\">A classic example of sexual selection involves the brightly coloured feathers of the peacock. The <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1471\"><strong>peacock<\/strong><\/a> is the male sex of the peafowl genera <em>Pavo<\/em> and <em>Afropavo. <\/em>During mating season, peacocks will fan their colourful tails wide and strut in front of the peahens in a grand display. The peahens will carefully observe these displays and will elect to mate with the male that they find the most appealing. Many studies have found that peahens prefer the males with the fullest, most colourful tails. While these large, showy tails provide a reproductive advantage, they can be a real burden in terms of escaping predators. The bright colours and patterns as well as the large size of the peacock tail make it difficult to hide. Once predators spot them, peacocks also struggle to fly away, with the heavy tail trailing behind and weighing them down (Figure 5.18). Some researchers have argued that the increased risk is part of the appeal for the peahens: only an especially strong, alert, and healthy peacock would be able to avoid predators while sporting such a spectacular tail.<\/p>\n<\/div>\n<p>It\u2019s important to keep in mind that sexual selection relies on the trait being present throughout mating years. Reflecting on the NF1 genetic disorder (see \u201cSpecial Topic: Neurofibromatosis Type 1 [NF1]\u201d), given how disfiguring the symptoms can become, some might find it surprising that half of the babies born with NF1 inherited it from a parent. Given that the disorder is autosomal dominant and fully penetrant (meaning it has no unaffected carriers), it may seem surprising that sexual selection doesn\u2019t exert more pressure against the mutated alleles. One important factor is that, while the neurofibromas typically begin to appear during puberty, they usually emerge only a few at a time and may grow very slowly. Many NF1 patients don\u2019t experience the more severe or disfiguring symptoms until later in life, long after they have started families of their own.<\/p>\n<p class=\"import-Normal\">Some researchers prefer to classify sexual selection separately, as a fifth force of evolution. The traits that underpin mate selection are entirely natural, of course. Research has shown that subtle traits, such as the type of pheromones (hormonal odors related to immune system alleles) someone emits and how those are perceived by the immune system genotype of the \u201csniffer,\u201d may play crucial and subconscious roles in whether we find someone attractive or not (Chaix, Cao, &amp; Donnelly 2008).<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Neurofibromatosis Type 1 (NF1)<\/h2>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1476\">Neurofibromatosis Type 1<\/a><\/strong>, also known as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1477\"><strong>NF1<\/strong><\/a>, is a genetic disorder that illustrates how a mutation in a single gene can affect multiple systems in the body. Surprisingly common, more people have NF1 than cystic fibrosis and muscular dystrophy combined. Even more surprising, given how common it is, is how few people have heard of it. One in every 3,000 babies is born with NF1, and this holds true for all populations worldwide (Riccardi 1992). This means that, for every 3,000 people in your community, there is likely at least one person living with this disorder. NF1 is an <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1478\">autosomal dominant<\/a> <\/strong>condition, which means that everyone born with a mutation in the gene, whether inherited or spontaneous, has a 50\/50 chance of passing it on to each of their own children.<\/p>\n<p class=\"import-Normal\">The NF1 disorder results from mutation of the <em>NF1<\/em> gene on Chromosome 17. Almost any mutation that affects the sequence of the gene\u2019s protein product, neurofibromin, will cause the disorder. Studies of individuals with NF1 have identified over 3,000 different mutations of all kinds (including point mutations, small and large indels, and translocations). The <em>NF1 <\/em>gene is one of the largest known genes, containing at least 60 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_724\">exons<\/a><\/strong> (protein-encoding sequences) in a span of about 300,000 nucleotides.<\/p>\n<p class=\"import-Normal\">We know that neurofibromin plays an important role in preventing tumor growth because one of the most common symptoms of the NF1 disorder is the growth of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1479\">benign<\/a> <\/strong>(noncancerous) tumors, called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1486\">neurofibromas<\/a><\/strong>. Neurofibromas sprout from nerve sheaths\u2014the tissues that encase our nerves\u2014throughout the body, usually beginning around puberty. There is no way to predict where the tumors will occur, or when or how quickly they will grow, although only about 15% turn <strong>malignant<\/strong> (cancerous). The two types of neurofibromas that are typically most visible are <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1480\"><strong>cutaneous neurofibromas<\/strong><\/a>, which are spherical bumps on, or just under, the surface of the skin (Figure 5.19), and <strong>plexiform neurofibromas<\/strong><em>, <\/em>growths involving whole branches of nerves, often giving the appearance that the surface of the skin is \u201cmelting\u201d (Figure 5.20).<\/p>\n<figure id=\"attachment_131\" aria-describedby=\"caption-attachment-131\" style=\"width: 510px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-129\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/4.18.jpg\" alt=\"A woman has dozens of round, skin-colored tumors visible on her face, neck, and hand.\" width=\"510\" height=\"340\" \/><figcaption id=\"caption-attachment-131\" class=\"wp-caption-text\">Figure 5.19: A woman with many cutaneous neurofibromas, a common symptom of Neurofibromatosis Type 1. Credit: <a class=\"rId105\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Woman with cutaneous neurofibromas (symptom of NF1)<\/a> by <a class=\"rId106\" href=\"https:\/\/positiveexposure.org\/about-the-program-2\/rick-guidotti\/\">Rick Guidotti of Positive Exposure<\/a> is used with permission and is available here under a <a class=\"rId107\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<figure id=\"attachment_131-2\" aria-describedby=\"caption-attachment-131-2\" style=\"width: 1900px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-130 size-full\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/4.19.jpg\" alt=\"An adult with large plexiform neurofibromas covering his face, none are on the child.\" width=\"1900\" height=\"700\" \/><figcaption id=\"caption-attachment-131-2\" class=\"wp-caption-text\">Figure 5.20: Photo on the left is of a man with large plexiform neurofibroma, another symptom of Neurofibromatosis Type 1. Photo on the right is a childhood photo of the same man, illustrating the progressive nature of the NF1 disorder. Credit: <a class=\"rId110\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Man with plexiform neurofibroma (symptom of NF1)<\/a> from Ashok Shrestha is used by permission and available here under a <a class=\"rId111\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. <a class=\"rId112\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Childhood photo of the same man with NF1 disorder<\/a> from Ashok Shrestha is used by permission and available here under a <a class=\"rId113\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Unfortunately, there is currently no cure for NF1. Surgical removal of neurofibromas risks paralysis, due to the high potential for nerve damage, and often results in the tumors growing back even more vigorously. This means that patients are often forced to live with disfiguring and often painful neurofibromas. People who are not familiar with NF1 often mistake neurofibromas for something contagious. This makes it especially hard for people living with NF1 to get jobs working with the public or even to enjoy spending time away from home. Raising public awareness about NF1 and its symptoms can be a great help in improving the quality of life for people living with this condition.<\/p>\n<figure style=\"width: 311px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image21-2.png\" alt=\"A child with darker oval birthmarks scattered across his torso and arms.\" width=\"311\" height=\"415\" \/><figcaption class=\"wp-caption-text\">Figure 5.21: Image of a child with caf\u00e9-au-lait macules (birthmarks) typical of the earliest symptoms of NF1. Credit: <a class=\"rId115\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Child with caf\u00e9-au-lait macules (birthmarks) typical of the earliest symptoms of NF1<\/a> by Andrea J. Alveshere is under a <a class=\"rId116\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">One of the first symptoms of NF1 in a small child is usually the appearance of <strong>caf\u00e9-au-lait spots<\/strong>, or <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1490\"><strong>CALS<\/strong><\/a>, which are flat, brown birthmark-like spots on the skin (Figure 5.21). CALS are often light brown, similar to the colour of coffee with cream, which is the reason for the name, although the shade of the pigment depends on a person\u2019s overall complexion. Some babies are born with CALS, but for others the spots appear within the first few years of life. Having six or more CALS larger than five millimetres (mm) across is a strong indicator that a child may have NF1.<\/p>\n<p class=\"import-Normal\">Other common symptoms include the following: gliomas (tumors) of the optic nerve, which can cause vision loss; thinning of bones and failure to heal if they break (often requiring amputation); low muscle tone (poor muscle development, often delaying milestones such as sitting up, crawling, and walking); hearing loss, due to neurofibromas on auditory nerves; and learning disabilities, especially those involving spatial reasoning. Approximately 50% of people with NF1 have some type of speech and\/or learning disability and often benefit greatly from early intervention services. Generalized developmental disability, however, is not common with NF1, so most people with NF1 live independently as adults. Many people with NF1 live full and successful lives, as long as their symptoms can be managed.<\/p>\n<p class=\"import-Normal\">Based on the wide variety of symptoms, it\u2019s clear that the neurofibromin protein plays important roles in many biochemical pathways. While everyone who has NF1 will exhibit some symptoms during their lifetime, there is a great deal of variation in the types and severity of symptoms, even between individuals from the same family who share the exact same NF1 mutation. It seems crazy that a gene with so many important functions would be so susceptible to mutation. Part of this undoubtedly has to do with its massive size\u2014a gene with 300,000 nucleotides has ten times more nucleotides available for mutation than does a gene of 30,000 bases. This also suggests that the mutability of this gene might provide some benefits, which is a possibility that we will revisit later in this chapter.<\/p>\n<\/div>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Sickle Cell Anemia<\/h2>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1492\">Sickle cell anemia<\/a><\/strong> is an autosomal recessive genetic disorder that affects millions of people worldwide. It is most common in Africa, countries around the Mediterranean Sea, and eastward as far as India. Populations in the Americas that have high percentages of ancestors from these regions also have high rates of sickle cell anemia. In the United States, it\u2019s estimated that 72,000 people live with the disease, with one in approximately 1,200 Hispanic-American babies and one in every 500 African-American babies inheriting the condition (World Health Organization 1996).<\/p>\n<figure style=\"width: 344px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image22-2.png\" alt=\"Round and sickle cells.\" width=\"344\" height=\"258\" \/><figcaption class=\"wp-caption-text\">Figure 5.22: Sickle cell anemia. Arrows indicate (a) sickled and (b) normal red blood cells. Credit: <a class=\"rId118\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Sickle-cell_smear_2015-09-10.jpg\">Sickle-cell smear 2015-09-10<\/a> by Paulo Henrique Orlandi Mourao has been modified (contrast modified and labels added) and is under a <a class=\"rId119\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Sickle cell anemia affects the hemoglobin protein in red blood cells. Normal red blood cells are somewhat doughnut-shaped\u2014round with a depression on both sides of the middle. They carry oxygen around the bloodstream to cells throughout the body. Red blood cells produced by the mutated form of the gene take on a stiff, sickle-like crescent shape when stressed by low oxygen or dehydration (Figure 5.22). Because of their elongated shape and the fact that they are stiff rather than flexible, they tend to form clumps in the blood vessels, inhibiting blood flow to adjacent areas of the body. This causes episodes of extreme pain and can cause serious problems in the oxygen-deprived tissues. The sickle cells also break down much more quickly than normal cells, often lasting only 20 days rather than the 120 days of normal cells. This causes an overall shortage of blood cells in the sickle cell patient, resulting in low iron (anemia) and problems associated with it such as extreme fatigue, shortness of breath, and hindrances to children\u2019s growth and development.<\/p>\n<p class=\"import-Normal\">The devastating effects of sickle cell anemia made its high frequency a pressing mystery. Why would an allele that is so deleterious in its homozygous form be maintained in a population at levels as high as the one in twelve African Americans estimated to carry at least one copy of the allele? The answer turned out to be one of the most interesting cases of balancing selection in the history of genetic study.<\/p>\n<p class=\"import-Normal\">While looking for an explanation, scientists noticed that the countries with high rates of sickle cell disease also shared a high risk for another disease called <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1493\"><strong>malaria<\/strong><\/a>, which is caused by infection of the blood by a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1494\"><strong><em>Plasmodium<\/em><\/strong><\/a> parasite. These parasites are carried by mosquitoes and enter the human bloodstream via a mosquito bite. Once infected, the person will experience flu-like symptoms that, if untreated, can often lead to death. Researchers discovered that many people living in these regions seemed to have a natural resistance to malaria. Further study revealed that people who carry the sickle cell allele are far less likely to experience a severe case of malaria. This would not be enough of a benefit to make the allele advantageous for the sickle cell homozygotes, who face shortened life spans due to sickle cell anemia. The real benefit of the sickle cell allele goes to the heterozygotes.<\/p>\n<p class=\"import-Normal\">People who are heterozygous for sickle cell carry one normal allele, which produces the normal, round, red blood cells, and one sickle cell allele, which produces the sickle-shaped red blood cells. Thus, they have both the sickle and round blood cell types in their bloodstream. They produce enough of the round red blood cells to avoid the symptoms of sickle cell anemia, but they have enough sickle cells to provide protection from malaria.<\/p>\n<p class=\"import-Normal\">When the <em>Plasmodium <\/em>parasites infect an individual, they begin to multiply in the liver, but then must infect the red blood cells to complete their reproductive cycle. When the parasites enter sickle-type cells, the cells respond by taking on the sickle shape. This prevents the parasite from circulating through the bloodstream and completing its life cycle, greatly inhibiting the severity of the infection in the sickle cell heterozygotes compared to non\u2013-sickle cell homozygotes. See Chapter 14 for more discussion of sickle cell anemia.<\/p>\n<\/div>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: The Real Primordial Cells\u2014<em>Dictyostelium Discoideum<\/em><\/h2>\n<p class=\"import-Normal\">The amoeba-like primordial cells that were used as recurring examples throughout this chapter are inspired by actual research that is truly fascinating. In 2015, Gareth Bloomfield and colleagues reported on their genomic study of the social amoeba <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1496\"><strong><em>Dictyostelium discoideum<\/em><\/strong> <\/a>(a.k.a. \u201cslime molds,\u201d although technically they are amoebae, not molds). Strains of these amoebae have been grown in research laboratories for many decades and are useful in studying the mechanisms that amoeboid single-celled organisms use to ingest food and liquid. For simplification of our examples in this chapter, our amoeba-like cells remained ocean dwellers. Wild <em>Dictyostelium discoideum<\/em>, however, live in soil and feed on soil bacteria by growing ruffles in their membranes that reach out to encapsulate the bacterial cell. Laboratory strains, however, are typically raised on liquid media (agar) in Petri dishes, which is not suitable for the wild-type amoebae. It was widely known that the laboratory strains must have developed mutations in one or more genes to allow them to ingest the larger nutrient particles in the agar and larger volumes of liquid, but the genes involved were not known.<\/p>\n<p class=\"import-Normal\">Bloomfield and colleagues performed genomic testing on both the wild and the laboratory strains of <em>Dictyostelium discoideum. <\/em>Their discovery was astounding: every one of the laboratory strains carried a mutation in the <em>NF1 <\/em>gene, the very same gene associated with Neurofibromatosis Type 1 (NF1) in humans. The antiquity of this massive, easily mutated gene is incredible. It originated in an ancestor common to both humans and these amoebae, and it has been retained in both lineages ever since. As seen in <em>Dictyostelium discoideum<\/em>, breaking the gene can be advantageous. Without a functioning copy of the neurofibromin protein, the cell membrane is able to form much-larger feeding structures, allowing the <em>NF1 <\/em>mutants to ingest larger particles and larger volumes of liquid. For these amoebae, this may provide dietary flexibility that functions somewhat like an insurance policy for times when the food supply is limited.<\/p>\n<p class=\"import-Normal\"><em>Dictyostelium discoideum <\/em>are also interesting in that they typically reproduce asexually, but under certain conditions, one cell will convert into a \u201cgiant\u201d cell, which encapsulates surrounding cells, transforming into one of three sexes. This cell will undergo meiosis, producing gametes that must combine with one of the other two sexes to produce viable offspring. This ability for sexual reproduction may be what allows <em>Dictyostelium discoideum<\/em> to benefit from the advantages of <em>NF1<\/em> mutation, while also being able to restore the wild type <em>NF1<\/em> gene in future generations.<\/p>\n<p class=\"import-Normal\">What does this mean for humans living with NF1? Well, understanding the role of the neurofibromin protein in the membranes of simple organisms like <em>Dictyostelium discoideum<\/em> may help us to better understand how it functions and malfunctions in the sheaths of human neurons. It\u2019s also possible that the mutability of the NF1 gene confers certain advantages to humans as well. Alleles of the NF1 gene have been found to reduce one\u2019s risk for alcoholism (Repunte-Canonigo Vez Et al. 2015), opiate addiction (Sanna Et al. 2002), Type 2 diabetes (Martins Et al. 2016), and hypomusicality (a lower-than-average musical aptitude; Cota Et al. 2018). This research is ongoing and will be exciting to follow in the coming years.<\/p>\n<\/div>\n<h2 class=\"__UNKNOWN__\">Studying Evolution in Action<\/h2>\n<div class=\"__UNKNOWN__\">\n<h3 class=\"import-Normal\"><strong>The Hardy-Weinberg Equilibrium <\/strong><\/h3>\n<p class=\"import-Normal\">This chapter has introduced you to the forces of evolution, the mechanisms by which evolution occurs. How do we detect and study evolution, though, in real time, as it happens? One tool we use is the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1497\"><strong>Hardy-<\/strong><strong>Weinberg<\/strong><strong> Equilibrium<\/strong><\/a>: a mathematical formula that allows estimation of the number and distribution of dominant and recessive alleles in a population. This aids in determining whether allele frequencies are changing and, if so, how quickly over time, and in favour of which allele? It\u2019s important to note that the Hardy-Weinberg formula only gives us an estimate based on the data for a snapshot in time. We will have to calculate it again later, after various intervals, to determine if our population is evolving and in what way the allele frequencies are changing.<\/p>\n<h3 class=\"import-Normal\">Calculating the Hardy-Weinberg Equilibrium<\/h3>\n<p class=\"import-Normal\">In the Hardy-Weinberg formula, <em>p <\/em>represents the frequency of the dominant allele, and <em>q<\/em> represents the frequency of the recessive allele. Remember, an allele\u2019s frequency is the proportion, or percentage, of that allele in the population. For the purposes of Hardy-Weinberg, we give the allele percentages as decimal numbers (e.g., 42% = 0.42), with the entire population (100% of alleles) equaling 1. If we can figure out the frequency of one of the alleles in the population, then it is simple to calculate the other. Simply subtract the known frequency from 1 (the entire population): 1<em> \u2013 p = q<\/em> and 1<em> \u2013 q = p<\/em>.<\/p>\n<p class=\"import-Normal\">The Hardy-Weinberg formula is <em>p<\/em><sup><em>2<\/em><\/sup><em> + 2pq + q<\/em><sup><em>2<\/em><\/sup>, where:<\/p>\n<p class=\"import-Normal\" style=\"padding-left: 40px\"><em>p<\/em><sup><em>2<\/em><\/sup> represents the frequency of the homozygous dominant genotype;<\/p>\n<p class=\"import-Normal\" style=\"padding-left: 40px\"><em>2pq<\/em> represents the frequency of the heterozygous genotype; and<\/p>\n<p class=\"import-Normal\" style=\"padding-left: 40px\"><em>q<\/em><sup><em>2<\/em><\/sup> represents the frequency of the homozygous recessive genotype.<\/p>\n<p class=\"import-Normal\">It is often easiest to determine <em>q<\/em><sup><em>2<\/em><\/sup> first, simply by counting the number of individuals with the unique, homozygous recessive phenotype (then dividing by the total individuals in the population to arrive at the \u201cfrequency\u201d). Once we have this number, we simply need to calculate the square root of the homozygous recessive phenotype frequency. That gives us <em>q.<\/em> Remember, 1 <em>\u2013<\/em> <em>q <\/em>equals <em>p<\/em>, so now we have the frequencies for both alleles in the population. If we needed to figure out the frequencies of heterozygotes and homozygous dominant genotypes, we\u2019d just need to plug the <em>p<\/em> and <em>q<\/em> frequencies back into the <em>p<\/em><sup><em>2<\/em><\/sup> and 2<em>pq<\/em> formulas.<\/p>\n<figure style=\"width: 329px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image25-1.png\" alt=\"A circle with seven grey and three white ladybugs.\" width=\"329\" height=\"347\" \/><figcaption class=\"wp-caption-text\">Figure 5.23: Ladybug population with a mixture of dark (red) and light (orange) individuals. Credit: <a class=\"rId129\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Ladybug mix (Figure 4.21)<\/a> original to <a class=\"rId130\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a class=\"rId131\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Let\u2019s imagine we have a population of ladybeetles that carries two alleles: a dominant allele that produces red ladybeetles and a recessive allele that produces orange ladybeetles. Since red is dominant, we\u2019ll use <em>R <\/em>to represent the red allele, and <em>r <\/em>to represent the orange allele. Our population has ten beetles, and seven are red and three are orange (Figure 5.23). Let\u2019s calculate the number of genotypes and alleles in this population.<\/p>\n<p class=\"import-Normal\">Of ten total beetles, we have three orange beetles3\/10 = .30 (30%) frequency\u2014and we know they are homozygous recessive (<em>rr<\/em>). So:<\/p>\n<p class=\"import-Normal\" style=\"text-indent: 36pt\"><em>rr = .3; <\/em>therefore, <em>r = <\/em>\u221a.3 = .5477<\/p>\n<p class=\"import-Normal\" style=\"text-indent: 36pt\"><em>R<\/em> = 1 <em>\u2013<\/em> .5477 = .4523<\/p>\n<p class=\"import-Normal\" style=\"text-indent: 36pt\">Using the Hardy-Weinberg formula:<\/p>\n<p class=\"import-Normal\" style=\"text-indent: 36pt\">1=.4523<sup>2 <\/sup>+ 2 x .4523 x .5477 +.5477<sup>2 <\/sup>= .20 + .50 + .30 = 1<\/p>\n<p class=\"import-Normal\" style=\"text-indent: 36pt\">Thus, the genotype breakdown is 20% <em>RR, <\/em>50%<em> Rr, <\/em>and 30%<em> rr <\/em><\/p>\n<p class=\"import-Normal\" style=\"text-indent: 36pt\">(2 red homozygotes, 5 red heterozygotes, and 3 orange homozygotes).<\/p>\n<p class=\"import-Normal\">Since we have 10 individuals, we know we have 20 total alleles: 4 red from the <em>RR<\/em> group, 5 red and 5 orange from the <em>Rr<\/em> group, and 6 orange from the <em>rr<\/em> group, for a grand total of 9 red and 11 orange (45% red and 55% orange, just like we estimated in the 1 \u2013 <em>q <\/em>step).<\/p>\n<p class=\"import-Normal\">Reminder: The Hardy-Weinberg formula only gives us an estimate for a snapshot in time. We will have to calculate it again later, after various intervals, to determine if our population is evolving and in what way the allele frequencies are changing.<\/p>\n<h3 class=\"import-Normal\"><strong>Interpreting Evolutionary Change: Nonra<\/strong><strong>ndom Mating <\/strong><\/h3>\n<p class=\"import-Normal\">Once we have detected change occurring in a population, we need to consider which evolutionary processes might be the cause of the change. It is important to watch for nonrandom mating patterns, to see if they can be included or excluded as possible sources of variation in allele frequencies.<\/p>\n<p class=\"import-Normal\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1498\"><strong>Nonrandom <\/strong><strong>m<\/strong><strong>ating<\/strong><\/a> (also known as assortative mating) occurs when mate choice within a population follows a nonrandom pattern.<\/p>\n<p class=\"import-Normal\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1499\"><strong>Positive assortative mating<\/strong><\/a> patterns result from a tendency for individuals to mate with others who share similar phenotypes. This often happens based on body size. Taking as an example dog breeds, it is easier for two Chihuahuas to mate and have healthy offspring than it is for a Chihuahua and a St. Bernard to do so. This is especially true if the Chihuahua is the female and would have to give birth to giant St. Bernard pups.<\/p>\n<p class=\"import-Normal\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1500\"><strong>Negative assortative mating<\/strong><\/a> patterns occur when individuals tend to select mates with qualities different from their own. This is what is at work when humans choose partners whose pheromones indicate that they have different and complementary immune alleles, providing potential offspring with a better chance at a stronger immune system.<\/p>\n<p class=\"import-Normal\">Among domestic animals, such as pets and livestock, assortative mating is often directed by humans who decide which pairs will mate to increase the chances of offspring having certain desirable traits. This is known as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1501\"><strong>a<\/strong><strong>rtificial <\/strong><strong>s<\/strong><strong>election<\/strong><em>.<\/em><\/a><\/p>\n<p class=\"import-Normal\">Among humans, in addition to phenotypic traits, cultural traits such as religion and ethnicity may also influence assortative mating patterns.<\/p>\n<h3 class=\"import-Normal\"><strong>Defining a Species<\/strong><\/h3>\n<p class=\"import-Normal\"><em>Species<\/em> are organisms whose individuals are capable of breeding because they are biologically and behaviourally compatible to produce viable, fertile offspring. <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1502\"><strong>Viable offspring<\/strong><\/a> are those offspring that are healthy enough to survive to adulthood. <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1503\"><strong>Fertile offspring<\/strong><\/a> are able to reproduce successfully, resulting in offspring of their own. Both conditions must be met for individuals to be considered part of the same species. As you can imagine, these criteria complicate the identification of distinct species in fossilized remains of extinct populations. In those cases, we must examine how much phenotypic variation is typically found within a comparable modern-day species; we can then determine whether the fossilized remains fall within the expected range of variation for a single species.<\/p>\n<p class=\"import-Normal\">Some species have subpopulations that are regionally distinct. These are classified as separate <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1505\">subspecies<\/a><\/strong> because they have their own unique phenotypes and are geographically isolated from one another. However, if they do happen to encounter one another, they are still capable of successful interbreeding.<\/p>\n<p class=\"import-Normal\">There are many examples of sterile hybrids that are offspring of parents from two different species. For example, horses and donkeys can breed and have offspring together. Depending on which species is the mother and which is the father, the offspring are either called mules, or hennies. Mules and hennies can live full life spans but are not able to have offspring of their own. Likewise, tigers and lions have been known to mate and have viable offspring. Again, depending on which species is the mother and which is the father, these offspring are called either ligers or tigons. Like mules and hennies, ligers and tigons are unable to reproduce. In each of these cases, the mismatched set of chromosomes that the offspring inherit produce an adequate set of functioning genes for the hybrid offspring; however, once mixed and divided in meiosis, the gametes don\u2019t contain the full complement of genes needed for survival in the third generation.<\/p>\n<h3 class=\"import-Normal\"><strong>Micro- to Macroevolution<\/strong><\/h3>\n<p class=\"import-Normal\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1506\"><strong>Microevolution<\/strong><\/a> refers to changes in allele frequencies within breeding populations\u2014that is, within single species. <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1507\"><strong>Macroevolution<\/strong><\/a> describes how the similarities and differences between species, as well as the phylogenetic relationships with other taxa, lead to changes that result in the emergence of new species. Consider our example of the peppered moth that illustrated microevolution over time, via directional selection favouring the peppered allele when the trees were clean and the dark pigment allele when the trees were sooty. Imagine that environmental regulations had cleaned up the air pollution in one part of the nation, while the coal-fired factories continued to spew soot in another area. If this went on long enough, it\u2019s possible that two distinct moth populations would eventually emerge\u2014one containing only the peppered allele and the other only harbouring the dark pigment allele.<\/p>\n<p class=\"import-Normal\">When a single population divides into two or more separate species, it is called <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1508\"><strong>speciation<\/strong><\/a>. The changes that prevent successful breeding between individuals who descended from the same ancestral population may involve chromosomal rearrangements, changes in the ability of the sperm from one species to permeate the egg membrane of the other species, or dramatic changes in hormonal schedules or mating behaviours that prevent members from the new species from being able to effectively pair up.<\/p>\n<p class=\"import-Normal\">There are two types of speciation: allopatric and sympatric. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1509\">Allopatric speciation<\/a><\/strong> is caused by long-term <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1510\"><strong>isolation<\/strong><\/a> (physical separation) of subgroups of the population (Figure 5.24). Something occurs in the environment\u2014perhaps a river changes its course and splits the group, preventing them from breeding with members on the opposite riverbank. Over many generations, new mutations and adaptations to the different environments on each side of the river may drive the two subpopulations to change so much that they can no longer produce fertile, viable offspring, even if the barrier is someday removed.<\/p>\n<figure style=\"width: 1000px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image23-2.png\" alt=\"Process of isolation leading to speciation, as described in the figure caption.\" width=\"1000\" height=\"379\" \/><figcaption class=\"wp-caption-text\">Figure 5.24: Isolation leading to speciation: a. original population before isolation; b. a barrier divides the population and prevents interbreeding between the two groups; c. time passes, and the populations become genetically distinct; d. after many generations, the two populations are no longer biologically or behaviourally compatible, thus can no longer interbreed, even if the barrier is removed. Credit: <a class=\"rId121\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Isolation Leading to Speciation (Figure 4.19)<\/a> original to <a class=\"rId122\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a class=\"rId123\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\"><strong>Sympatric speciation<\/strong> occurs when the population splits into two or more separate species while remaining located together <em>without<\/em> a physical barrier. This typically results from a new mutation that pops up among some members of the population that prevents them from successfully reproducing with anyone who does not carry the same mutation. This is seen particularly often in plants, as they have a higher frequency of chromosomal duplications.<\/p>\n<p class=\"import-Normal\">One of the quickest rates of speciation is observed in the case of adaptive radiation. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1512\">Adaptive radiation<\/a><\/strong> refers to the situation in which subgroups of a single species rapidly diversify and adapt to fill a variety of ecological niches. An <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1513\"><strong>e<\/strong><strong>cological niche<\/strong><\/a> is a set of constraints and resources that is available in an environmental setting. Evidence for adaptive radiations is often seen after population bottlenecks. A mass disaster kills off many species, and the survivors have access to a new set of territories and resources that were either unavailable or much coveted and fought over before the disaster. The offspring of the surviving population will often split into multiple species, each of which stems from members in that first group of survivors who happened to carry alleles that were advantageous for a particular niche.<\/p>\n<p class=\"import-Normal\">The classic example of adaptive radiation brings us back to Charles Darwin and his observations of the many species of finches on the Galapagos Islands. We are still not sure how the ancestral population of finches first arrived on that remote Pacific Island chain, but they found themselves in an environment filled with various insects, large and tiny seeds, fruit, and delicious varieties of cactus. Some members of that initial population carried alleles that gave them advantages for each of these dietary niches. In subsequent generations, others developed new mutations, some of which were beneficial. These traits were selected for, making the advantageous alleles more common among their offspring. As the finches spread from one island to the next, they would be far more likely to find mates among the birds on their new island. Birds feeding in the same area were then more likely to mate together than birds who have different diets, contributing to additional assortative mating. Together, these evolutionary mechanisms caused rapid speciation that allowed the new species to make the most of the various dietary niches (Figure 5.25).<\/p>\n<figure style=\"width: 619px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image24-1.png\" alt=\"A family tree of finches with different sized beaks.\" width=\"619\" height=\"325\" \/><figcaption class=\"wp-caption-text\">Figure 5.25: Darwin\u2019s finches demonstrating Adaptive Radiation. Credit: <a class=\"rId125\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Darwin\u2019s finches (Figure 4.20)<\/a> original to <a class=\"rId126\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a class=\"rId127\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">In today\u2019s modern world, understanding these evolutionary processes is crucial for developing immunizations and antibiotics that can keep up with the rapid mutation rate of viruses and bacteria. This is also relevant to our food supply, which relies, in large part, on the development of herbicides and pesticides that keep up with the mutation rates of pests and weeds. Viruses, bacteria, agricultural pests, and weeds have all shown great flexibility in developing alleles that make them resistant to the latest medical treatment, pesticide, or herbicide. Billion-dollar industries have specialized in trying to keep our species one step ahead of the next mutation in the pests and infectious diseases that put our survival at risk.<\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Review Questions<strong><br \/>\n<\/strong><\/h2>\n<ul>\n<li>Summarize the Modern Synthesis and provide several examples of how it is relevant to questions and problems in our world today.<\/li>\n<li>You inherit a house from a long-lost relative that contains a fancy aquarium, filled with a variety of snails. The phenotypes include large snails and small snails; red, black, and yellow snails; and solid, striped, and spotted snails. Devise a series of experiments that would help you determine how many snail species are present in your aquarium.<\/li>\n<li>Match the correct force of evolution with the correct real-world example:<br \/>\na. Mutationi. 5-alpha reductase deficiency<br \/>\nb. Genetic Driftii. Peppered Moths<br \/>\nc. Gene Flowiii. Neurofibromatosis Type 1<br \/>\nd. Natural Selectioniv. Scutellata Honey Bees<\/li>\n<li>Imagine a population of common house mice (<em>Mus musculus<\/em>). Draw a comic strip illustrating how mutation, genetic drift, gene flow, and natural selection might transform this population over several (or more) generations.<\/li>\n<li>\n<p class=\"import-Normal\">The many breeds of the single species of domestic dog (<em>Canis<\/em> <em>familiaris<\/em>) provide an extreme example of microevolution. Discuss why this is the case. What future scenarios can you imagine that could potentially transform the domestic dog into an example of macroevolution?<\/p>\n<\/li>\n<li>\n<p class=\"import-Normal\">The ability to roll one\u2019s tongue (lift the outer edges of the tongue to touch each other, forming a tube) is a dominant trait. In a small town of 1,500 people, 500 can roll their tongues. Use the Hardy-Weinberg formula to determine how many individuals in the town are homozygous dominant, heterozygous, and homozygous recessive.<\/p>\n<\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\">Key Terms<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\"><strong>5-alpha reductase deficiency<\/strong>: An autosomal recessive syndrome that manifests when a child having both X and Y sex chromosomes inherits two nonfunctional (mutated) copies of the SRD5A2 gene, producing a deficiency in a hormone necessary for development in infancy of typical male genitalia. These children often appear at birth to have female genitalia, but they develop a penis and other sexual characteristics when other hormones kick in during puberty.<\/p>\n<p class=\"import-Normal\"><strong>Adaptive radiation<\/strong>: The situation in which subgroups of a single species rapidly diversify and adapt to fill a variety of ecological niches.<\/p>\n<p class=\"import-Normal\"><strong>Admixture<\/strong>: A term often used to describe gene flow between human populations. Sometimes also used to describe gene flow between nonhuman populations.<\/p>\n<p class=\"import-Normal\"><strong>Allele frequency<\/strong>: The ratio, or percentage, of one allele compared to the other alleles for that gene within the study population.<\/p>\n<p class=\"import-Normal\"><strong>Alleles<\/strong>: Variant forms of genes.<\/p>\n<p class=\"import-Normal\"><strong>Allopatric speciation<\/strong>: Speciation caused by long-term isolation (physical separation) of subgroups of the population.<\/p>\n<p class=\"import-Normal\"><strong>Antibiotics<\/strong>: Medicines prescribed to treat bacterial infections.<\/p>\n<p class=\"import-Normal\"><strong>Artificial selection<\/strong>: Human-directed assortative mating among domestic animals, such as pets and livestock, designed to increase the chances of offspring having certain desirable traits.<\/p>\n<p class=\"import-Normal\"><strong>Asexual reproduction<\/strong>: Reproduction via mitosis, whereby offspring are clones of the parents.<\/p>\n<p class=\"import-Normal\"><strong>Autosomal dominant<\/strong>: A phenotype produced by a gene on an autosomal chromosome that is expressed, to the exclusion of the recessive phenotype, in heterozygotes.<\/p>\n<p class=\"import-Normal\"><strong>Autosomal recessive<\/strong>: A phenotype produced by a gene on an autosomal chromosome that is expressed only in individuals homozygous for the recessive allele.<\/p>\n<p class=\"import-Normal\"><strong>Balanced translocations<\/strong>: Chromosomal translocations in which the genes are swapped but no genetic information is lost.<\/p>\n<p class=\"import-Normal\"><strong>Balancing selection<\/strong>: A pattern of natural selection that occurs when the extremes of a trait are selected against, favouring the intermediate phenotype (a.k.a. stabilizing selection).<\/p>\n<p class=\"import-Normal\"><strong>Beneficial mutations<\/strong>: Mutations that produce some sort of an advantage to the individual.<\/p>\n<p class=\"import-Normal\"><strong>Benign<\/strong>: Noncancerous. Benign tumors may cause problems due to the area in which they are located (e.g., they might put pressure on a nerve or brain area), but they will not release cells that aggressively spread to other areas of the body.<\/p>\n<p class=\"import-Normal\"><strong>Caf\u00e9-au-lait spots (CALS)<\/strong>: Flat, brown birthmark-like spots on the skin, commonly associated with Neurofibromatosis Type 1.<\/p>\n<p class=\"import-Normal\"><strong>Chromosomal translocations<\/strong>: The transfer of DNA between nonhomologous chromosomes.<\/p>\n<p class=\"import-Normal\"><strong>Chromosomes<\/strong>: Molecules that carry collections of genes.<\/p>\n<p class=\"import-Normal\"><strong>Codons<\/strong>: Three-nucleotide units of DNA that function as three-letter \u201cwords,\u201d encoding instructions for the addition of one amino acid to a protein or indicating that the protein is complete.<\/p>\n<p class=\"import-Normal\"><strong>Cretaceous\u2013Paleogene extinction<\/strong>: A mass disaster caused by an asteroid that struck the earth approximately 66 million years ago and killed 75% of life on Earth, including all terrestrial dinosaurs. (a.k.a. K-Pg Extinction, Cretatious-Tertiary Extinction, and K-T Extinction).<\/p>\n<p class=\"import-Normal\"><strong>Crossover events<\/strong>: Chromosomal alterations that occur when DNA is swapped between homologous chromosomes while they are paired up during meiosis I.<\/p>\n<p class=\"import-Normal\"><strong>Cutaneous neurofibromas<\/strong>: Neurofibromas that manifest as spherical bumps on or just under the surface of the skin.<\/p>\n<p class=\"import-Normal\"><strong>Deleterious mutation<\/strong>: A mutation producing negative effects to the individual such as the beginnings of cancers or heritable disorders.<\/p>\n<p class=\"import-Normal\"><strong>Deletions<\/strong>: Mutations that involve the removal of one or more nucleotides from a DNA sequence.<\/p>\n<p class=\"import-Normal\"><strong>Derivative chromosomes<\/strong>: New chromosomal structures resulting from translocations.<\/p>\n<p class=\"import-Normal\"><strong><em>Dictyostelium discoideum<\/em><\/strong>: A species of social amoebae that has been widely used for laboratory research. Laboratory strains of <em>Dictyostelium discoideum <\/em>all carry mutations in the <em>NF1<\/em> gene, which is what allows them to survive on liquid media (agar) in Petri dishes.<\/p>\n<p class=\"import-Normal\"><strong>Directional selection<\/strong>: A pattern of natural selection in which one phenotype is favoured over the other, causing the frequencies of the associated advantageous alleles to gradually increase.<\/p>\n<p class=\"import-Normal\"><strong>Disruptive selection<\/strong>: A pattern of natural selection that occurs when both extremes of a trait are advantageous and intermediate phenotypes are selected against (a.k.a. diversifying selection).<\/p>\n<p class=\"import-Normal\"><strong>DNA repair mechanisms<\/strong>: Enzymes that patrol and repair DNA in living cells.<\/p>\n<p class=\"import-Normal\"><strong>DNA transposons<\/strong>: Transposons that are clipped out of the DNA sequence itself and inserted elsewhere in the genome.<\/p>\n<p class=\"import-Normal\"><strong>Ecological niche<\/strong>: A set of constraints and resources that are available in an environmental setting.<\/p>\n<p class=\"import-Normal\"><strong>Ellis-van Creveld syndrome<\/strong>: An autosomal recessive disorder characterized by short stature (dwarfism), polydactyly (the development of more than five digits [fingers or toes] on the hands or feet), abnormal tooth development, and heart defects. Estimated to affect approximately one in 60,000 individuals worldwide, among the Old Order Amish of Lancaster County, the rate is estimated to be as high as one in every 200 births.<\/p>\n<p class=\"import-Normal\"><strong>Evolution<\/strong>: A change in the allele frequencies in a population over time.<\/p>\n<p class=\"import-Normal\"><strong>Exons<\/strong>: The DNA sequences within a gene that directly encode protein sequences. After being transcribed into messenger RNA, the introns (DNA sequences within a gene that do not directly encode protein sequences) are clipped out, and the exons are pasted together prior to translation.<\/p>\n<p class=\"import-Normal\"><strong>Fertile offspring<\/strong>: Offspring that can successfully reproduce, resulting in offspring of their own.<\/p>\n<p class=\"import-Normal\"><strong>Founder effect<\/strong>: A type of genetic drift that occurs when members of a population leave the main or \u201cparent\u201d group and form a new population that no longer interbreeds with the other members of the original group.<\/p>\n<p class=\"import-Normal\"><strong>Frameshift mutations<\/strong>: Types of indels that involve the insertion or deletion of any number of nucleotides that is not a multiple of three. These \u201cshift the reading frame\u201d and cause all codons beyond the mutation to be misread.<\/p>\n<p class=\"import-Normal\"><strong>Gametes<\/strong>: The reproductive cells, produced through meiosis (a.k.a. germ cells or sperm or egg cells).<\/p>\n<p class=\"import-Normal\"><strong>Gene<\/strong>: A sequence of DNA that provides coding information for the construction of proteins.<\/p>\n<p class=\"import-Normal\"><strong>Gene flow<\/strong>: The movement of alleles from one population to another. This is one of the forces of evolution.<\/p>\n<p class=\"import-Normal\"><strong>Gene pool<\/strong>: The entire collection of genetic material in a breeding community that can be passed on from one generation to the next.<\/p>\n<p class=\"import-Normal\"><strong>Genetic drift<\/strong>: Random changes in allele frequencies within a population from one generation to the next. This is one of the forces of evolution.<\/p>\n<p class=\"import-Normal\"><strong>Genotype<\/strong>: The set of alleles that an individual has for a given gene.<\/p>\n<p class=\"import-Normal\"><strong>Genotype frequencies<\/strong>: The ratios or percentages of the different homozygous and heterozygous genotypes in the population.<\/p>\n<p class=\"import-Normal\"><strong><em>Guevedoces<\/em><\/strong>: The term coined locally in the Dominican Republic for the condition scientifically known as 5-alpha reductase deficiency. The literal translation is \u201cpenis at twelve.\u201d<\/p>\n<p class=\"import-Normal\"><strong>Hardy-Weinberg Equilibrium<\/strong>: A mathematical formula (<em>1=p<\/em><sup><em>2<\/em><\/sup><em> + 2pq + q<\/em><sup><em>2<\/em><\/sup> ) that allows estimation of the number and distribution of dominant and recessive alleles in a population.<\/p>\n<p class=\"import-Normal\"><strong>Harlequin ladybeetle<\/strong>: A species of ladybeetle, native to East Asia, that was introduced to Europe and the Americas as a form of pest control. After many decades of use, one of the North American strains developed the ability to reproduce in diverse environments, causing it to spread rapidly throughout the Americas, Europe, and Africa. It has hybridized with European strains and is now a major pest in its own right.<\/p>\n<p class=\"import-Normal\"><strong>Heterozygous genotype<\/strong>: A genotype comprising two different alleles.<\/p>\n<p class=\"import-Normal\"><strong>Homozygous genotype<\/strong>: A genotype comprising an identical set of alleles.<\/p>\n<p class=\"import-Normal\"><strong>Hybridization<\/strong>: A term often used to describe gene flow between nonhuman populations.<\/p>\n<p class=\"import-Normal\"><strong>Inbreeding<\/strong>: The selection of mates exclusively from within a small, closed population.<\/p>\n<p class=\"import-Normal\"><strong>Indels<\/strong>: A class of mutations that includes both insertions and deletions.<\/p>\n<p class=\"import-Normal\"><strong>Inherited mutation<\/strong>: A mutation that has been passed from parent to offspring.<\/p>\n<p class=\"import-Normal\"><strong>Insertions<\/strong>: Mutations that involve the addition of one or more nucleotides into a DNA sequence.<\/p>\n<p class=\"import-Normal\"><strong>Isolation<\/strong>: Prevention of a population subgroup from breeding with other members of the same species due to a physical barrier or, in humans, a cultural rule.<\/p>\n<p class=\"import-Normal\"><strong>Last Universal Common Ancestor (LUCA)<\/strong>: The ancient organism from which all living things on Earth are descended.<\/p>\n<p class=\"import-Normal\"><strong>Macroevolution<\/strong>: Changes that result in the emergence of new species, how the similarities and differences between species, as well as the phylogenetic relationships with other taxa, lead to changes that result in the emergence of new species.<\/p>\n<p class=\"import-Normal\"><strong>Malaria<\/strong>: A frequently deadly mosquito-borne disease caused by infection of the blood by a <em>Plasmodium<\/em> parasite.<\/p>\n<p class=\"import-Normal\"><strong>Malignant<\/strong>: Cancerous. Malignant tumors grow aggressively and their cells may metastasize (travel through the blood or lymph systems) to form new, aggressive tumors in other areas of the body.<\/p>\n<p class=\"import-Normal\"><strong>Microevolution<\/strong>: Changes in allele frequencies within breeding populations\u2014that is, within a single species.<\/p>\n<p class=\"import-Normal\"><strong>Modern Synthesis<\/strong>: The integration of Darwin\u2019s, Mendel\u2019s, and subsequent research into a unified theory of evolution.<\/p>\n<p class=\"import-Normal\"><strong>Monosomies<\/strong>: Conditions resulting from a nondisjunction event, in which a cell ends up with only one copy of a chromosome. In humans, a single X chromosome is the only survivable monosomy.<\/p>\n<p class=\"import-Normal\"><strong>Mutation<\/strong>: A change in the nucleotide sequence of the genetic code. This is one of the forces of evolution.<\/p>\n<p class=\"import-Normal\"><strong>Natural selection<\/strong>: An evolutionary process that occurs when certain phenotypes confer an advantage or disadvantage in survival and\/or reproductive success. This is one of the forces of evolution, and it was first identified by Charles Darwin.<\/p>\n<p class=\"import-Normal\"><strong>Negative assortative mating<\/strong>: A pattern that occurs when individuals tend to select mates with qualities different from their own.<\/p>\n<p class=\"import-Normal\"><strong>Neurofibromas<\/strong>: Nerve sheath tumors that are common symptoms of Neurofibromatosis Type 1.<\/p>\n<p class=\"import-Normal\"><strong>Neurofibromatosis Type 1<\/strong>: An autosomal dominant genetic disorder affecting one in every 3,000 people. It is caused by mutation of the <em>NF1<\/em> gene on Chromosome 17, resulting in a defective neurofibromin protein. The disorder is characterized by neurofibromas, caf\u00e9-au-lait spots, and a host of other potential symptoms.<\/p>\n<p class=\"import-Normal\"><strong>NF1<\/strong>: An abbreviation for Neurofibromatosis Type 1. When italicized, <em>NF1 <\/em>refers to the gene on Chromosome 17 that encodes the neurofibromin protein.<\/p>\n<p class=\"import-Normal\"><strong>Nondisjunction events<\/strong>: Chromosomal abnormalities that occur when the homologous chromosomes (in meiosis I) or sister chromatids (in meiosis II and mitosis) fail to separate after pairing. The result is that both chromosomes or chromatids end up in the same daughter cell, leaving the other daughter cell without any copy of that chromosome.<\/p>\n<p class=\"import-Normal\"><strong>Nonrandom mating<\/strong>: A scenario in which mate choice within a population follows a nonrandom pattern (a.k.a. assortative mating).<\/p>\n<p class=\"import-Normal\"><strong>Nonsynonymous mutation<\/strong>: A point mutation that causes a change in the resulting protein.<\/p>\n<p class=\"import-Normal\"><strong>Old Order Amish<\/strong>: A culturally isolated population in Lancaster County, Pennsylvania, that has approximately 50,000 current members, all of whom can trace their ancestry back to a group of approximately eighty individuals. This group has high rates of certain genetics disorders, including Ellis-van Creveld syndrome.<\/p>\n<p class=\"import-Normal\"><strong>Origins of life<\/strong>: How the first living organism came into being.<\/p>\n<p class=\"import-Normal\"><strong>Peacock<\/strong>: The male sex of the peafowl, famous for its large, colourful tail, which it dramatically displays to attract mates. (The female of the species is known as a peahen.)<\/p>\n<p class=\"import-Normal\"><strong>Peppered moth<\/strong>: A species of moth (<em>Biston betularia<\/em>) found in England that has light and dark phenotypes. During the Industrial Revolution, when soot blackened the trees, the frequency of the previously rare dark phenotype dramatically increased, as lighter-coloured moths were easier for birds to spot against the sooty trees. After environmental regulations eliminated the soot, the lighter-coloured phenotype gradually became most common again.<\/p>\n<p class=\"import-Normal\"><strong>Phenotype<\/strong>: The observable traits that are produced by a genotype.<\/p>\n<p class=\"import-Normal\"><strong>Phylogenetic tree of life<\/strong>: A family tree of all living organisms, based on genetic relationships.<\/p>\n<p class=\"import-Normal\"><strong>Phylogenies<\/strong>: Genetically determined family lineages.<\/p>\n<p class=\"import-Normal\"><strong><em>Plasmodium<\/em><\/strong>: A genus of mosquito-borne parasite. Several <em>Plasmodium<\/em> species cause malaria when introduced to the human bloodstream via a mosquito bite.<\/p>\n<p class=\"import-Normal\"><strong>Plexiform neurofibromas<\/strong>: Neurofibromas that involve whole branches of nerves, often giving the appearance that the surface of the skin is \u201cmelting.\u201d<\/p>\n<p class=\"import-Normal\"><strong>Point mutation<\/strong>: A single-letter (single-nucleotide) change in the genetic code, resulting in the substitution of one nucleic acid base for a different one.<\/p>\n<p class=\"import-Normal\"><strong>Polymorphisms<\/strong>: Multiple forms of a trait; alternative phenotypes within a given species.<\/p>\n<p class=\"import-Normal\"><strong>Population<\/strong>: A group of individuals who are genetically similar enough and geographically near enough to one another that they can breed and produce new generations of individuals.<\/p>\n<p class=\"import-Normal\"><strong>Population bottleneck<\/strong>: A type of genetic drift that occurs when the number of individuals in a population drops dramatically due to some random event.<\/p>\n<p class=\"import-Normal\"><strong>Positive assortative mating<\/strong>: A pattern that results from a tendency for individuals to mate with others who share similar phenotypes.<\/p>\n<p class=\"import-Normal\"><strong>Retrotransposons<\/strong>: Transposons that are transcribed from DNA into RNA, and then are \u201creverse transcribed,\u201d to insert the copied sequence into a new location in the DNA.<\/p>\n<p class=\"import-Normal\"><strong>Scutellata honey bees<\/strong>: A strain of honey bees that resulted from the hybridization of African and European honey bee subspecies. These bees were accidentally released into the wild in 1957 in Brazil and have since spread throughout South and Central America and into the United States. Also known as \u201ckiller bees,\u201d they tend to be very aggressive in defence of their hives and have caused many fatal injuries to humans and livestock.<\/p>\n<p class=\"import-Normal\"><strong>Sexual reproduction<\/strong>: Reproduction via meiosis and combination of gametes. Offspring inherit genetic material from both parents.<\/p>\n<p class=\"import-Normal\"><strong>Sexual selection<\/strong>: An aspect of natural selection in which the selective pressure specifically affects reproductive success (the ability to successfully breed and raise offspring).<\/p>\n<p class=\"import-Normal\"><strong>Sickle cell anemia<\/strong>: An autosomal recessive genetic disorder that affects millions of people worldwide. It is most common in Africa, countries around the Mediterranean Sea, and eastward as far as India. Homozygotes for the recessive allele develop the disorder, which produce misshapen red blood cells that cause iron deficiency, painful episodes of oxygen-deprivation in localized tissues, and a host of other symptoms. In heterozygotes, though, the sickle cell allele confers a greater resistance to malaria.<\/p>\n<p class=\"import-Normal\"><strong>Somatic cells<\/strong>: The cells of our organs and other body tissues (all cells except gametes) that replicate by mitosis.<\/p>\n<p class=\"import-Normal\"><strong>Speciation<\/strong>: The process by which a single population divides into two or more separate species.<\/p>\n<p class=\"import-Normal\"><strong>Species<\/strong>: Organisms whose individuals are capable of breeding because they are biologically and behaviourally compatible to produce viable, fertile offspring.<\/p>\n<p class=\"import-Normal\"><strong>Spontaneous mutation<\/strong>: A mutation that occurs due to random chance or unintentional exposure to mutagens. In families, a spontaneous mutation is the first case, as opposed to mutations that are inherited from parents.<\/p>\n<p class=\"import-Normal\"><strong>Subspecies<\/strong>: A distinct subtype of a species. Most often, this is a geographically isolated population with unique phenotypes; however, it remains biologically and behaviourally capable of interbreeding with other populations of the same species.<\/p>\n<p class=\"import-Normal\"><strong>Sympatric speciation<\/strong>: When a population splits into two or more separate species while remaining located together without a physical (or cultural) barrier.<\/p>\n<p class=\"import-Normal\"><strong>Synonymous mutation<\/strong>: A point mutation that does not change the resulting protein.<\/p>\n<p class=\"import-Normal\"><strong>Transposable elements<\/strong>: Fragments of DNA that can \u201cjump\u201d around in the genome.<\/p>\n<p class=\"import-Normal\"><strong>Transposon<\/strong>: Another term for \u201ctransposable element.\u201d<\/p>\n<p class=\"import-Normal\"><strong>Trisomies<\/strong>: Conditions in which three copies of the same chromosome end up in a cell, resulting from a nondisjunction event. Down syndrome, Edwards syndrome, and Patau syndrome are trisomies.<\/p>\n<p class=\"import-Normal\"><strong>Unbalanced translocations<\/strong>: Chromosomal translocations in which there is an unequal exchange of genetic material, resulting in duplication or loss of genes.<\/p>\n<p class=\"import-Normal\"><strong>UV crosslinking<\/strong>: A type of mutation in which adjacent thymine bases bind to one another in the presence of UV light.<\/p>\n<p class=\"import-Normal\"><strong>Viable offspring<\/strong>: Offspring that are healthy enough to survive to adulthood.<\/p>\n<p class=\"import-Normal\"><strong>Xeroderma pigmentosum<\/strong>: An autosomal recessive disease in which DNA repair mechanisms do not function correctly, resulting in a host of problems especially related to sun exposure, including severe sunburns, dry skin, heavy freckling, and other pigment changes.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration<\/h2>\n<p>Explore Evolution on <a href=\"https:\/\/www.hhmi.org\/biointeractive\/evolution-collection\">HHMI\u2019s Biointeractive website<\/a>.<\/p>\n<p>Teaching Evolution through <a href=\"https:\/\/humanorigins.si.edu\/education\/teaching-evolution-through-human-examples\">Human Examples, Smithsonian Museum of Natural History websites<\/a>.<\/p>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\">Bloomfield, Gareth, David Traynor, Sophia P. Sander, Douwe M. Veltman, Justin A. Pachebat, and Robert R. Kay. 2015. \u201cNeurofibromin Controls Macropinocytosis and Phagocytosis in <em>Dictyostelium<\/em>.\u201d <em>eLife<\/em> 4:e04940.<\/p>\n<p class=\"import-Normal\">Chaix, Rapha\u00eblle, Chen Cao, and Peter Donnelly. 2008. \u201cIs Mate Choice in Humans MHC-Dependent?\u201d <em>PLoS Genetics<\/em> 4 (9): e1000184.<\/p>\n<p class=\"import-Normal\">Cook, Laurence M. 2003. \"The Rise and Fall of the <em>Carbonaria<\/em> Form of the Peppered Moth.\" <em>The Quarterly Review of Biology<\/em> 78 (4): 399\u2013417.<\/p>\n<p class=\"import-Normal\">Cota, Bruno C\u00e9zar Lage, Jo\u00e3o Gabriel Marques Fonseca, Luiz Oswaldo Carneiro Rodrigues, Nilton Alves de Rezende, Pollyanna Barros Batista, Vincent Michael Riccardi, and Luciana Macedo de Resende. 2018. \u201cAmusia and Its Electrophysiological Correlates in Neurofibromatosis Type 1.\u201d <em>Arquivos de Neuro-Psiquiatria<\/em> 76 (5): 287\u2013295.<\/p>\n<p class=\"import-Normal\">D\u2019Asdia, Maria Cecilia, Isabella Torrente, Federica Consoli, Rosangela Ferese, Monia Magliozzi, Laura Bernardini, Valentina Guida, et al. 2013. \u201cNovel and Recurrent EVC and EVC2 Mutations in Ellis-van Creveld Syndrome and Weyers Acrofacial Dyostosis.\u201d <em>European Journal of Medical Genetics<\/em> 56 (2): 80\u201387.<\/p>\n<p class=\"import-Normal\">Dobzhansky, Theodosius. 1937. <em>Genetics and the Origin of Species. <\/em>Columbia University Biological Series. New York: Columbia University Press.<\/p>\n<p class=\"import-Normal\">Facon, Beno\u00eet, Laurent Crespin, Anne Loiseau, Eric Lombaert, Alexandra Magro, and Arnaud Estoup. 2011. \u201cCan Things Get Worse When an Invasive Species Hybridizes? The Harlequin Ladybird <em>Harmonia axyridis<\/em> in France as a Case Study.\u201d <em>Evolutionary Applications<\/em> 4 (1): 71\u201388.<\/p>\n<p class=\"import-Normal\">Fisher, Ronald A. 1919. \"The Correlation between Relatives on the Supposition of Mendelian Inheritance.\" <em>Transactions of the Royal Society of Edinburgh<\/em> 52 (2): 399\u2013433.<\/p>\n<p class=\"import-Normal\">Ford, E. B. 1942. <em>Genetics for Medical Students<\/em>. London: Methuen.<\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff\">Ford, E. B. 1949. <em>Mendelism and Evolution<\/em>. London: Methuen.<\/p>\n<p class=\"import-Normal\">Grant, Bruce S. 1999. \u201cFine-tuning the Peppered Moth Paradigm.\u201d <em>Evolution<\/em> 53 (3): 980\u2013984.<\/p>\n<p class=\"import-Normal\">Haldane, J. B. S. 1924. \u201cA Mathematical Theory of Natural and Artificial Selection (Part 1).\u201d <em>Transactions of the Cambridge Philosophical Society<\/em> 23 (2):19\u201341.<\/p>\n<p>Hoelzel, A. R., Gkafas, G. A., Kang, H., Sarigol, F., Le Boeuf, B., Costa, D. P., Beltran, R. S., Reiter, J., Robinson, P. W., McInerney, N., Seim, I., Sun, S., Fan, G., &amp; Li, S. (2024). Genomics of post-bottleneck recovery in the northern elephant seal. Nature Ecology &amp; Evolution, 8, 686\u2013694. https:\/\/doi.org\/10.1038\/s41559-024-02337-4<\/p>\n<p class=\"import-Normal\">Imperato-McGinley, J., and Y.-S. Zhu. 2002. \u201cAndrogens and Male Physiology: The Syndrome of 5 Alpha-Reductase-2 Deficiency.\u201d <em>Molecular and Cellular Endocrinology <\/em>198 (1-2): 51\u201359.<\/p>\n<p class=\"import-Normal\">Jablonski, David, and W. G. Chaloner. 1994. \"Extinctions in the Fossil Record.\u201d <em>Philosophical Transactions of the Royal Society of London B: Biological Sciences<\/em> 344 (1307): 11\u201317.<\/p>\n<p class=\"import-Normal\">Livi-Bacci, Massimo. 2006. \u201cThe Depopulation of Hispanic America after the Conquest.\u201d <em>Population Development and Review<\/em> 32 (2): 199\u2013232.<\/p>\n<p class=\"import-Normal\">Lombaert, Eric, Thomas Guillemaud, Jean-Marie Cornuet, Thibaut Malausa, Beno\u00eet Facon, and Arnaud Estoup. 2010. \"Bridgehead Effect in the Worldwide Invasion of the Biocontrol Harlequin Ladybird.\u201d <em>PLoS ONE<\/em> 5 (3): e9743.<\/p>\n<p class=\"import-Normal\">Martins, Aline Stangherlin, Ann Kristine Jansen, Luiz Oswaldo Carneiro Rodrigues, Camila Maria Matos, Marcio Leandro Ribeiro Souza, Juliana Ferreira de Souza, Maria de F\u00e1tima Haueisen Sander Diniz, et al. 2016. \u201cLower Fasting Blood Glucose in Neurofibromatosis Type 1.\u201d <em>Endocrine Connections<\/em> 5 (1): 28\u201333.<\/p>\n<p class=\"import-Normal\">Pickering, Gary, James Lin, Roland Riesen, Andrew Reynolds, Ian Brindle, and George Soleas. 2004. \"Influence of <em>Harmonia axyridis<\/em> on the Sensory Properties of White and Red Wine.\" <em>American Journal of Enology and Viticulture<\/em> 55 (2): 153\u2013159.<\/p>\n<p class=\"import-Normal\">Repunte-Canonigo Vez, Melissa A. Herman, Tomoya Kawamura, Henry R. Kranzler, Richard Sherva, Joel Gelernter, Lindsay A. Farrer, Marisa Roberto, and Pietro Paolo Sanna. 2015. \u201cNF1 Regulates Alcohol Dependence-Associated Excessive Drinking and Gamma-Aminobutyric Acid Release in the Central Amygdala in Mice and Is Associated with Alcohol Dependence in Humans.\u201d <em>Biological Psychiatry<\/em> 77 (10): 870\u2013879.<\/p>\n<p class=\"import-Normal\">Riccardi, Vincent M. 1992. <em>Neurofibromatosis: Phenotype, Natural History, and Pathogenesis.<\/em> Baltimore: Johns Hopkins University Press.<\/p>\n<p class=\"import-Normal\">Sanford, Malcolm T. 2006. \"The Africanized Honey Bee in the Americas: A Biological Revolution with Human Cultural Implications, Part V\u2014Conclusion.\" <em>American Bee Journal <\/em>146 (7): 597\u2013599.<\/p>\n<p class=\"import-Normal\">Sanna, Pietro Paolo, Cindy Simpson, Robert Lutjens, and George Koob. 2002. \u201cERK Regulation in Chronic Ethanol Exposure and Withdrawal.\u201d <em>Brain Research<\/em> 948 (1\u20132): 186\u2013191.<\/p>\n<p>Weber, DianaS., Stewart, B. S., Garza, J. Carlos., &amp; Lehman, N. (2000). An empirical genetic assessment of the severity of the northern elephant seal population bottleneck. Current Biology, 10(20), 1287\u20131290. https:\/\/doi.org\/10.1016\/s0960-9822(00)00759-4<\/p>\n<p class=\"import-Normal\">World Health Organization. 1996. \u201cControl of Hereditary Disorders: Report of WHO Scientific meeting (1996).\u201d WHO Technical Reports 865. Geneva: World Health Organization.<\/p>\n<p class=\"import-Normal\">World Health Organization. 2017. \u201cGlobal Priority List of Antibiotic-Resistant Bacteria to Guide Research, Discovery, and Development of New Antibiotics.\u201d Global Priority Pathogens List, February 27. Geneva: World Health Organization. https:\/\/www.who.int\/medicines\/publications\/WHO-PPL-Short_Summary_25Feb-ET_NM_WHO.pdf.<\/p>\n<p class=\"import-Normal\">Wright, Sewall. 1932. \"The Roles of Mutation, Inbreeding, Crossbreeding, and Selection in Evolution.\" <em>Proceedings of the Sixth International Congress on Genetics<\/em> 1 (6): 356\u2013366.<\/p>\n<h2 class=\"import-Normal\">Acknowledgement<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\">Many thanks to Dr. Vincent M. Riccardi for sharing his vast knowledge of neurofibromatosis and for encouraging me to explore it from an anthropological perspective.<\/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_1683_800\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_800\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<p class=\"import-Normal\">Andrea J. Alveshere, Ph.D., Western Illinois University<\/p>\n<h6>Student contributors for this chapter: Corin Laberge, Hazel Moorcroft, Isabella Michel, Julian J. Pantoja Quiroz<\/h6>\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__-3\/\"><em>Chapter 4: Forces of Evolution<\/em><\/a><em>\u201d by Andrea J. Alveshere. 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: #000000\">Learning Objectives<\/span><\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li class=\"import-Normal\">Outline a 21st-century perspective of the Modern Synthesis.<\/li>\n<li class=\"import-Normal\">Define populations and population genetics as well as the methods used to study them.<\/li>\n<li class=\"import-Normal\">Identify the forces of evolution and become familiar with examples of each.<\/li>\n<li class=\"import-Normal\">Discuss the evolutionary significance of mutation, genetic drift, gene flow, and natural selection.<\/li>\n<li class=\"import-Normal\">Explain how allele frequencies can be used to study evolution as it happens.<\/li>\n<li class=\"import-Normal\">Contrast micro- and macroevolution.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p>It\u2019s hard for us, with our typical human life spans of less than 100 years, to imagine all the way back, 3.8 billion years ago, to the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1442\">origins of life<\/a><\/strong>. Scientists still study and debate how life came into being and whether it originated on Earth or in some other region of the universe (including some scientists who believe that studying evolution can reveal the complex processes that were set in motion by God or a higher power). What we do know is that a living single-celled organism was present on Earth during the early stages of our planet\u2019s existence. This organism had the potential to reproduce by making copies of itself, just like bacteria, many amoebae, and our own living cells today. In fact, with modern technologies, we can now trace genetic lineages, or <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1290\">phylogenies<\/a><\/strong>, and determine the relationships between all of today\u2019s living organisms\u2014eukaryotes (animals, plants, fungi, etc.), archaea, and bacteria\u2014on the branches of the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1444\">phylogenetic tree of life<\/a><\/strong> (Figure 5.1).<\/p>\n<figure style=\"width: 675px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2023\/02\/image1-1.png\" alt=\"Branches lead off of a single celled universal ancestor to images of bacteria, archaea, and eukarya (represented by a mouse, mushroom, and fern, among others).\" width=\"675\" height=\"475\" \/><figcaption class=\"wp-caption-text\">Figure 5.1: Phylogenetic tree of life illustrating probable relationships between the single-celled Last Universal Common Ancestor (LUCA) and select examples of bacteria, archaea, and eukaryotes. Major evolutionary developments, including independent evolution of multicellularity, photosynthesis, and respiration, are indicated along the branches. <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 class=\"rId11\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Cladograma_dos_Dom%C3%ADnios_e_Reinos.png\">Cladograma dos Dominios e Reinos<\/a> by <a class=\"rId12\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:MarceloTeles\">MarceloTeles<\/a> has been modified (English labels replace Portuguese) and is under a <a class=\"rId13\" 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\">Looking at the common sequences in modern genomes, we can even make educated guesses about the likely genetic sequence of the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1446\"><strong>Last Universal Common Ancestor (LUCA)<\/strong><\/a> of all living things. Through a wondrous series of mechanisms and events over nearly four billion years, that ancient single-celled organism gave rise to the rich diversity of species that fill the lands, seas, and skies of our planet. This chapter explores the mechanisms by which that amazing transformation occurred and considers some of the crucial scientific experiments that shaped our current understanding of the evolutionary process.<\/p>\n<h2 class=\"import-Normal\">Population Genetics<\/h2>\n<h3 class=\"import-Normal\"><strong>Defining Populations and the Variations <\/strong><strong>w<\/strong><strong>ithin Them<\/strong><\/h3>\n<p class=\"import-Normal\">One of the major breakthroughs in understanding the mechanisms of evolutionary change came with the realization that evolution takes place at the level of populations, not within individuals. In the biological sciences, a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1454\"><strong>p<\/strong><strong>opulation<\/strong><\/a> is defined as a group of individuals of the same <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1456\"><strong>species<\/strong><\/a> who are geographically near enough to one another that they can breed and produce new generations of individuals.<\/p>\n<p class=\"import-Normal\">For the purpose of studying evolution, we recognize populations by their even smaller units: genes. Remember, a\u00a0<a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1458\"><strong>gene<\/strong><\/a> is the basic unit of information that encodes the proteins needed to grow and function as a living organism. Each gene can have multiple <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_738\">alleles<\/a><\/strong>, or variants\u2014each of which may produce a slightly different protein. Each individual, for genetic inheritance purposes, carries a collection of genes that can be passed down to future generations. For this reason, in population genetics, we think of populations as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1462\"><strong>gene pools<\/strong><\/a>, which refers to the entire collection of genetic material in a breeding community that can be passed on from one generation to the next.<\/p>\n<p class=\"import-Normal\">For genes carried on our human chromosomes (our nuclear DNA), we inherit two copies of each, one from each parent. This means we may carry two of the same alleles (a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1464\"><strong>homozygous genotype<\/strong><\/a>) or two different alleles (a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1466\"><strong>heterozygous<\/strong> <strong>genotype<\/strong><\/a>) for each nuclear gene.<\/p>\n<h3 class=\"import-Normal\"><strong>Defining Evolution <\/strong><\/h3>\n<p class=\"import-Normal\">In order to understand evolution, it\u2019s crucial to remember that evolution is always studied at the population level. Also, if a population were to stay exactly the same from one generation to the next, it would not be evolving. So evolution requires both a population of breeding individuals and some kind of a genetic change occurring within it. Thus, the simple definition of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1468\"><strong>evolution<\/strong><\/a> is a change in the allele frequencies in a population over time. What do we mean by allele frequencies? <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1470\"><strong>Allele frequencies<\/strong> <\/a>refer to the ratio, or percentage, of one allele (one variant of a gene) compared to the other alleles for that gene within the study population (Figure 5.2). By contrast, <strong>genotype frequencies<\/strong> are the ratios or percentages of the different homozygous and heterozygous genotypes in the population. Because we carry two alleles per <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_736\">genotype<\/a><\/strong>, the total count of alleles in a population will usually be exactly double the total count of genotypes in the same population (with the exception being rare cases in which an individual carries a different number of chromosomes than the typical two; e.g., Down syndrome results when a child carries three copies of Chromosome 21).<\/p>\n<figure style=\"width: 652px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image2.jpg\" alt=\"Genotypes are represented as combinations of alleles and allele frequencies.\" width=\"652\" height=\"883\" \/><figcaption class=\"wp-caption-text\">Figure 5.2: Population evolution can be measured by allele frequency changes. This diagram illustrates the differences between genotype frequencies and allele frequencies, as well as how they can be measured in a population of snapdragon flowers. The lower portion of the diagram also depicts how evolution is recognized as allele frequencies change in a population over time. <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>.\u00a0Credit: Population evolution original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Katie Nelson and Beth Shook is a collective work under a <a class=\"rId15\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\">CC BY-NC 4.0 License<\/a>. [Includes <a class=\"rId16\" href=\"https:\/\/pixabay.com\/vectors\/snapdragon-flower-pink-lilac-plant-146850\/\">Snapdragon-flower-pink-lilac<\/a> by <a class=\"rId17\" href=\"https:\/\/pixabay.com\/users\/openclipart-vectors-30363\/\">OpenClipart-Vectors<\/a>, <a class=\"rId18\" href=\"https:\/\/creativecommons.org\/share-your-work\/public-domain\/cc0\/\">public domain (CC0)<\/a> under a <a class=\"rId19\" href=\"https:\/\/pixabay.com\/service\/terms\/\">Pixabay License<\/a>.]<\/figcaption><\/figure>\n<h2 class=\"import-Normal\">The Forces of Evolution<\/h2>\n<p class=\"import-Normal\">Today, we recognize that evolution takes place through a combination of mechanisms: mutation, genetic drift, gene flow, and natural selection. These mechanisms are called the \u201cforces of evolution\u201d; together they account for all the genotypic variation observed in the world today. Keep in mind that each of these forces was first defined and then tested\u2014and retested\u2014through the experimental work of the many scientists who contributed to the Modern Synthesis.<\/p>\n<h3 class=\"import-Normal\"><strong>Mutation<\/strong><\/h3>\n<p class=\"import-Normal\">The first force of evolution we will discuss is mutation, and for good reason: mutation is the original source of all the genetic variation found in every living thing. Imagine all the way back in time to the very first single-celled organism, floating in Earth\u2019s primordial sea. Based on what we observe in simple, single-celled organisms today, that organism probably spent its lifetime absorbing nutrients and dividing to produce cloned copies of itself. While the numbers of individuals in that population would have grown (as long as the environment was favorable), nothing would have changed in that perfectly cloned population. There would not have been variety among the individuals. It was only through a copying error\u2014the introduction of a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1474\"><strong>mutation<\/strong><\/a>, or change, into the genetic code\u2014that new alleles were introduced into the population.<br style=\"clear: both\" \/><br style=\"clear: both\" \/>After many generations have passed in our primordial population, mutations have created distinct chromosomes. The cells are now amoeba-like, larger than many of their tiny bacterial neighbors, who have long since become their favorite source of nutrients. Without mutation to create this diversity, all living things would still be identical to LUCA, our universal ancestor (Figure 5.3).<\/p>\n<figure style=\"width: 663px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image3-2.png\" alt=\"Universal Ancestor linked to the Eukarya branch.\" width=\"663\" height=\"338\" \/><figcaption class=\"wp-caption-text\">Figure 5.3: Key mutational differences between Last Universal Common Ancestor and an amoeba-like primordial cell. <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<strong>: <\/strong>Key differences between LUCA and a primordial cell original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Andrea J. Alveshere is a collective work under a <a class=\"rId21\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Includes <a class=\"rId22\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Cladograma_dos_Dom%C3%ADnios_e_Reinos.png\">Cladograma dos Dominios e Reinos<\/a> by <a class=\"rId23\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:MarceloTeles\">MarceloTeles<\/a> (cropped, labels and color changed), <a class=\"rId24\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0<\/a><a class=\"rId25\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">; <\/a><a class=\"rId26\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Amoeba_proteus_TK-UT.svg\">Amoeba Proteus TK-UT<\/a> by <a class=\"rId27\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Nefronus\">Tom\u00e1\u0161 Kebert<\/a> and <a class=\"rId28\" href=\"https:\/\/www.umimeto.org\/\">umimeto.org<\/a> (cropped and color changed), <a class=\"rId29\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0<\/a>.]<\/figcaption><\/figure>\n<p class=\"import-Normal\">When we think of genetic mutation, we often first think of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1476\">deleterious mutations<\/a><\/strong>\u2014the ones associated with negative effects such as the beginnings of cancers or heritable disorders. The fact is, though, that every genetic adaptation that has helped our ancestors survive since the dawn of life is directly due to <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1478\">beneficial mutations<\/a><\/strong>\u2014changes in the DNA that provided some sort of advantage to a given population at a particular moment in time. For example, a beneficial mutation allowed chihuahuas and other tropical-adapted dog breeds to have much thinner fur coats than their cold-adapted cousins the northern wolves, malamutes, and huskies.<\/p>\n<figure style=\"width: 320px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-1-1.png\" alt=\"UV radiation damages nucleotides in DNA.\" width=\"320\" height=\"248\" \/><figcaption class=\"wp-caption-text\">Figure 5.4: A crosslinking mutation in which a UV photon induces a bond between two thymine bases. <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<strong>: <\/strong><a class=\"rId31\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">UV-induced Thymine dimer mutation (Figure 4.6)<\/a> original to <a class=\"rId32\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a class=\"rId33\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Every one of us has genetic mutations. Yes, even you. The DNA in some of your cells today differs from the original DNA that you inherited when you were a tiny, fertilized egg. Mutations occur all the time in the cells of our skin and other organs, due to chemical changes in the nucleotides. Exposure to the UV radiation in sunlight is one common cause of skin mutations. Interaction with UV light causes <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1480\"><strong>UV crosslinking<\/strong><\/a>, in which adjacent thymine bases bind with one another (Figure 5.4). Many of these mutations are detected and corrected by <strong>DNA repair mechanisms<\/strong>, enzymes that patrol and repair DNA in living cells, while other mutations may cause a new freckle or mole or, perhaps, an unusual hair to grow. For people with the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1484\">autosomal recessive<\/a><\/strong> disease <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1486\"><strong>xeroderma pigmentosum<\/strong><\/a>, these repair mechanisms do not function correctly, resulting in a host of problems especially related to sun exposure, including severe sunburns, dry skin, heavy freckling, and other pigment changes.<\/p>\n<p class=\"import-Normal\">Most of our mutations exist in <strong>somatic<\/strong> cells, which are the cells of our organs and other body tissues. Those will not be passed onto future generations and so will not affect the population over time. Only mutations that occur in the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_686\">gametes<\/a><\/strong>, the reproductive cells (i.e., the sperm or egg cells), will be passed onto future generations. When a new mutation pops up at random in a family lineage, it is known as a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1490\"><strong>spontaneous mutation<\/strong><\/a>. If the individual born with this spontaneous mutation passes it on to his offspring, those offspring receive an <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1492\">inherited mutation<\/a><\/strong>. Geneticists have identified many classes of mutations and the causes and effects of many of these.<\/p>\n<h4 class=\"import-Normal\"><em>Point Mutations<\/em><\/h4>\n<p class=\"import-Normal\">A <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1494\">point mutation<\/a><\/strong> is a single-letter (single-nucleotide) change in the genetic code resulting in the substitution of one nucleic acid base for a different one. As you learned in Chapter 3, the DNA code in each gene is translated through three-letter \u201cwords\u201d known as <strong>codons<\/strong>. So depending on how the point mutation changes the \u201cword,\u201d the effect it will have on the protein may be major or minor or may make no difference at all.<\/p>\n<p class=\"import-Normal\">If a mutation does not change the resulting protein, then it is called a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1496\">synonymous mutation<\/a><\/strong>. Synonymous mutations do involve a letter (nucleic acid) change, but that change results in a codon that codes for the same \u201cinstruction\u201d (the same amino acid or stop code) as the original codon. Mutations that do cause a change in the protein are known as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1498\"><strong>nonsynonymous mutations<\/strong>.<\/a> Nonsynonymous mutations may change the resulting protein\u2019s amino acid sequence by altering the DNA sequence that encodes the mRNA or by changing how the mRNA is spliced prior to translation (refer to Chapter 3 for more details).<\/p>\n<h4 class=\"import-Normal\"><em>Insertions and Deletions<\/em><\/h4>\n<p class=\"import-Normal\">In addition to point mutations, another class of mutations are <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1500\"><strong>insertions<\/strong><\/a> and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1504\">deletions<\/a><\/strong>, or <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1502\"><strong>indels<\/strong><\/a>, for short. As the name suggests, these involve the addition (insertion) or removal (deletion) of one or more coding sequence letters (nucleic acids). These typically first occur as an error in DNA replication, wherein one or more nucleotides are either duplicated or skipped in error. Entire codons or sets of codons may also be removed or added if the indel is a multiple of three nucleotides.<\/p>\n<p class=\"import-Normal\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1506\"><strong>Frameshift<\/strong> <strong>mutations<\/strong><\/a> are types of indels that involve the insertion or deletion of any number of nucleotides that is not a multiple of three (e.g., adding one or two extra letters to the code). Because these indels are not consistent with the codon numbering, they \u201cshift the reading frame,\u201d causing all the codons beyond the mutation to be misread. Like point mutations, small indels can also disrupt splice sites.<\/p>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1508\">Transposable elements<\/a><\/strong>, or <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1510\">transposons<\/a><\/strong>, are fragments of DNA that can \u201cjump\u201d around in the genome. There are two types of transposons: <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1512\">retrotransposons<\/a><\/strong> are transcribed from DNA into RNA and then \u201creverse transcribed,\u201d to insert the copied sequence into a new location in the DNA, and<strong> <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1514\">DNA transposons<\/a><\/strong>, which do not involve RNA. DNA transposons are clipped out of the DNA sequence itself and inserted elsewhere in the genome. Because transposable elements insert themselves into existing DNA sequences, they are frequent gene disruptors. At certain times, and in certain species, it appears that transposons became very active, likely accelerating the mutation rate (and thus, the genetic variation) in those populations during the active periods.<\/p>\n<h4 class=\"import-Normal\"><em>Chromosomal Alterations <\/em><\/h4>\n<p class=\"import-Normal\">The final major category of genetic mutations are changes at the chromosome level: crossover events, nondisjunction events, and translocations. <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1516\"><strong>Crossover events<\/strong> <\/a> occur when DNA is swapped between homologous chromosomes while they are paired up during meiosis I. Crossovers are thought to be so common that some DNA swapping may happen every time chromosomes go through meiosis I. Crossovers don\u2019t necessarily introduce new alleles into a population, but they do make it possible for new combinations of alleles to exist on a single chromosome that can be passed to future generations. This also enables new combinations of alleles to be found within siblings who share the same parents. Also, if the fragments that cross over don\u2019t break at exactly the same point, they can cause genes to be deleted from one of the homologous chromosomes and duplicated on the other.<\/p>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1518\">Nondisjunction events<\/a><\/strong> occur when the homologous chromosomes (in meiosis I) or sister chromatids (in meiosis II and mitosis) fail to separate after pairing. The result is that both chromosomes or chromatids end up in the same daughter cell, leaving the other daughter cell without any copy of that chromosome (Figure 5.5). Most nondisjunctions at the gamete level are fatal to the embryo. The most widely known exception is Trisomy 21, or Down syndrome, which results when an embryo inherits three copies of Chromosome 21: two from one parent (due to a nondisjunction event) and one from the other (Figure 5.6). <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1520\">Trisomies<\/a> <\/strong>(triple chromosome conditions) of Chromosomes 18 (Edwards syndrome) and 13 (Patau syndrome) are also known to result in live births, but the children usually have severe complications and rarely survive beyond the first year of life.<\/p>\n<figure style=\"width: 601px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5.jpg\" alt=\"Egg cell undergoes normal meiosis and nondisjunction in meisosis 1.\" width=\"601\" height=\"391\" \/><figcaption class=\"wp-caption-text\">Figure 5.5: Illustration of an egg cell (oocyte) undergoing normal meiosis 1, resulting in a diploid daughter cell, compared to an egg cell undergoing nondisjunction during meiosis 1, resulting in a trisomy in the daughter cell. Credit: <a class=\"rId35\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Trisomy_due_to_nondisjunction_in_maternal_meiosis_1.png\">Trisomy due to nondisjunction in maternal meiosis 1<\/a> by Wpeissner has been modified (labels deleted by Katie Nelson) and is under a <a class=\"rId36\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<figure style=\"width: 316px\" class=\"wp-caption alignright\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image6-1.jpg\" alt=\"A young woman in a blue polo shirt smiles at the camera.\" width=\"316\" height=\"364\" \/><figcaption class=\"wp-caption-text\">Figure 5.6: Amy Bockerstette, a competitive golfer and disabilities advocate, also has Down Syndrome. Credit: <a class=\"rId38\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Amy_Bockerstette_Headshot.jpg\">Amy Bockerstette Headshot<\/a> by Bucksgrandson is under a <a class=\"rId39\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Sex chromosome trisomies (XXX, XXY, XYY) and X chromosome <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1522\"><strong>monosomies<\/strong> <\/a>(inheritance of an X chromosome from one parent and no sex chromosome from the other) are also survivable and fairly common. The symptoms vary but often include atypical sexual characteristics, either at birth or at puberty, and often result in sterility. The X chromosome carries unique genes that are required for survival; therefore, Y chromosome monosomies are incompatible with life.<\/p>\n<p class=\"import-Normal\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1524\"><strong>Chromosomal translocations<\/strong><\/a> involve transfers of DNA between nonhomologous chromosomes. This may involve swapping large portions of two or more chromosomes. The exchanges of DNA may be balanced or unbalanced. In <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1526\"><strong>balanced translocations<\/strong><\/a>, the genes are swapped, but no genetic information is lost. In <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1528\">unbalanced translocations<\/a><\/strong>, there is an unequal exchange of genetic material, resulting in duplication or loss of genes. Translocations result in new chromosomal structures called <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1530\"><strong>derivative chromosomes<\/strong><\/a>, because they are derived or created from two different chromosomes<em>. <\/em>Translocations are often found to be linked to cancers and can also cause infertility. Even if the translocations are balanced in the parent, the embryo often won\u2019t survive unless the baby inherits both of that parent\u2019s derivative chromosomes (to maintain the balance).<\/p>\n<h3 class=\"import-Normal\"><strong>Genetic Drift<\/strong><\/h3>\n<p class=\"import-Normal\">The second force of evolution is commonly known as genetic drift. This is an unfortunate misnomer, as this force actually involves the drifting of alleles, not genes. <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1532\"><strong>Genetic <\/strong><strong>d<\/strong><strong>rift<\/strong><\/a> refers to <em>random<\/em> changes (\u201cdrift\u201d) in allele frequencies from one generation to the next. The genes are remaining constant within the population; it is only the alleles of the genes that are changing in frequency. The random nature of genetic drift is a crucial point to understand: it specifically occurs when none of the variant alleles confer an advantage.<\/p>\n<figure style=\"width: 368px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image7-2.png\" alt=\"A smooth cell has a gently curving exterior surface, and a ruffled cell has undulating surface.\" width=\"368\" height=\"215\" \/><figcaption class=\"wp-caption-text\">Figure 5.7: Smooth and ruffled amoeba-like cells. Credit: Smooth and ruffled amoeba-like cells original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Andrea J. Alveshere is a collective work under a <a class=\"rId41\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Includes <a class=\"rId42\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Amoeba_proteus_TK-UT.svg\">Amoeba Proteus TK-UT<\/a> by <a class=\"rId43\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Nefronus\">Tom\u00e1\u0161 Kebert<\/a> and <a class=\"rId44\" href=\"https:\/\/www.umimeto.org\/\">umimeto.org<\/a> (modified), <a class=\"rId45\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0<\/a>.]<\/figcaption><\/figure>\n<p class=\"import-Normal\">Let\u2019s imagine far back in time, again, to that ancient population of amoeba-like cells, subsisting and occasionally dividing, in the primordial sea. A mutation occurs in one of the cells that changes the texture of the cell membrane from a relatively smooth surface to a highly ruffled one (Figure 5.7). This has absolutely no effect on the cell\u2019s quality of life or ability to reproduce. In fact, eyes haven\u2019t evolved yet, so no one in the world at the time would even notice the difference. The cells in the population continue to divide, and the offspring of the ruffled cell inherit the ruffled membrane. The frequency (percentage) of the ruffled allele in the population, from one generation to the next, will depend entirely on how many offspring that first ruffled cell ends up having, and the random events that might make the ruffled alleles more common or more rare (such as population bottlenecks and founder effects, which are discussed below).<\/p>\n<h4 class=\"import-Normal\"><em>Sexual Reproduction and Random Inheritance<\/em><\/h4>\n<p class=\"import-Normal\">Tracking alleles gets a bit more complicated in our primordial cells when, after a number of generations, a series of mutations have created populations that reproduce sexually. These cells now must go through an extra round of cell division (meiosis) to create haploid gametes. The combination of two gametes is now required to produce each new diploid offspring.<\/p>\n<figure style=\"width: 262px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image8-1.png\" alt=\"A Punnett square with ruffled and smooth cells.\" width=\"262\" height=\"262\" \/><figcaption class=\"wp-caption-text\">Figure 5.8: A Punnett square demonstrating the sexual inheritance pattern of ruffled (dominant) and smooth amoeba-like primordial cells. Credit: Punnett square of primordial cells original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Andrea J. Alveshere is a collective work under a <a class=\"rId47\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Includes <a class=\"rId48\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Amoeba_proteus_TK-UT.svg\">Amoeba Proteus TK-UT<\/a> by <a class=\"rId49\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Nefronus\">Tom\u00e1\u0161 Kebert<\/a> and <a class=\"rId50\" href=\"https:\/\/www.umimeto.org\/\">umimeto.org<\/a> (modified), <a class=\"rId51\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0<\/a>; <a class=\"rId52\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Punnett_hetero_x_hetero.svg\">Punnett Hetero x Hetero<\/a> by <a class=\"rId53\" href=\"https:\/\/commons.wikimedia.org\/w\/index.php?title=User:Purpy_Pupple&amp;redirect=no\">Purpy Pupple<\/a> (modified), <a class=\"rId54\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/deed.en\">CC BY-SA 3.0<\/a>].<\/figcaption><\/figure>\n<p class=\"import-Normal\">In the earlier population, which reproduced via <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1534\">asexual reproduction<\/a><\/strong>, a cell either carried the smooth allele or the ruffled allele. With <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1536\">sexual reproduction<\/a><\/strong>, a cell inherits one allele from each parent, so there are homozygous cells that contain two smooth alleles, homozygous cells that contain two ruffled alleles, and heterozygous cells that contain one of each allele (Figure 5.8). If the new, ruffled allele happens to be dominant (and we\u2019ll imagine that it is), the heterozygotes will have ruffled cell <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1238\"><strong>phenotypes<\/strong><\/a> but also will have a 50\/50 chance of passing on a smooth allele to each offspring. As long as neither phenotype (ruffled nor smooth) provides any advantage over the other, the variation in the population from one generation to the next will remain completely random.<\/p>\n<p class=\"import-Normal\">In sexually reproducing populations (including humans and many other animals and plants in the world today), that 50\/50 chance of inheriting one or the other allele from each parent plays a major role in the random nature of genetic drift.<\/p>\n<h4 class=\"import-Normal\"><em>Population Bottlenecks <\/em><\/h4>\n<p class=\"import-Normal\">A <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1538\"><strong>population bottleneck<\/strong><\/a> occurs when the number of individuals in a population drops dramatically due to some random event. The most obvious, familiar examples are natural disasters. Tsunamis and hurricanes devastating island and coastal populations and forest fires and river floods wiping out populations in other areas are all too familiar. When a large portion of a population is randomly wiped out, the allele frequencies (i.e., the percentages of each allele) in the small population of survivors are often much different from the frequencies in the predisaster, or \u201cparent,\u201d population.<\/p>\n<p class=\"import-Normal\">If such an event happened to our primordial ocean cell population\u2014perhaps a volcanic fissure erupted in the ocean floor and only the cells that happened to be farthest from the spewing lava and boiling water survived\u2014we might end up, by random chance, with a surviving population that had mostly ruffled alleles, in contrast to the parent population, which had only a small percentage of ruffles (Figure 5.9).<\/p>\n<figure style=\"width: 665px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image9-2.png\" alt=\"Ruffled and smooth cells experience population bottleneck when a lava flow divides the populations.\" width=\"665\" height=\"332\" \/><figcaption class=\"wp-caption-text\">Figure 5.9: Illustration of a population of amoeba-like cells shifting from primarily smooth phenotypes (at left) to mostly ruffled phenotypes due to eruption of a volcanic fissure (at right) that exterminated the nearest cells. Credit: Population of amoeba-like cells and volcanic fissure original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Andrea J. Alveshere is a collective work under a <a class=\"rId56\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Includes <a class=\"rId57\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Amoeba_proteus_TK-UT.svg\">Amoeba Proteus TK-UT<\/a> by <a class=\"rId58\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Nefronus\">Tom\u00e1\u0161 Kebert<\/a> and <a class=\"rId59\" href=\"https:\/\/www.umimeto.org\/\">umimeto.org<\/a> (modified), <a class=\"rId60\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0<\/a>.]<\/figcaption><\/figure>\n<p class=\"import-Normal\">One of the most famous examples of a population bottleneck is the prehistoric disaster that led to the extinction of dinosaurs, the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1540\"><strong>Cretaceous\u2013Paleogene <\/strong><strong>extinction<\/strong><\/a> event (often abbreviated K\u2013Pg; previously K-T). This occurred approximately 66 million years ago. Dinosaurs and all their neighbors were going about their ordinary routines when a massive asteroid zoomed in from space and crashed into what is now the Gulf of Mexico, creating an impact so enormous that populations within hundreds of miles of the crash site were likely immediately wiped out. The skies filled with dust and debris, causing temperatures to plummet worldwide. It\u2019s estimated that 75% of the world\u2019s species went extinct as a result of the impact and the deep freeze that followed (Jablonski and Chaloner 1994).<\/p>\n<figure style=\"width: 399px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image10-2.png\" alt=\" A rat-like creature sits atop a dinosaur, raising a fist in a victorious gesture.\" width=\"399\" height=\"323\" \/><figcaption class=\"wp-caption-text\">Figure 5.10: The Cretaceous\u2013Paleogene extinction event, which led to the fall of the dinosaurs and rise of the mammals. Credit: <a class=\"rId62\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">The<\/a> <a class=\"rId64\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Cretaceous\u2013Paleogene extinction event (Figure 4.12)<\/a> original to <a class=\"rId65\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is 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\">The populations that emerged from the K-Pg extinction were markedly different from their pre-disaster communities. Surviving mammal populations expanded and diversified, and other new creatures appeared. The ecosystems of Earth were filled with new organisms and have never been the same (Figure 5.10).<\/p>\n<p class=\"import-Normal\">Much more recently in geological time, during the colonial period, many human populations experienced bottlenecks as a result of the fact that imperial powers were inclined to slaughter communities who were reluctant to give up their lands and resources. This effect was especially profound in the Americas, where Indigenous populations faced the compounded effects of brutal warfare, exposure to new bacteria and viruses (against which they had no immunity), and ultimately segregation on resource-starved reservations. The populations in Europe, Asia, and Africa had experienced regular gene flow during the 10,000-year period in which most kinds of livestock were being domesticated, giving them many generations of experience building up immunity against zoonotic diseases (those that can pass from animals to humans). In contrast, the residents of the Americas had been almost completely isolated during those millennia, so all these diseases swept through the Americas in rapid succession, creating a major loss of genetic diversity in the Indigenous American population. It is estimated that between 50% and 95% of the Indigenous American populations died during the first decades after European contact, around 500 years ago (Livi-Bacci 2006).<\/p>\n<p class=\"import-Normal\">An urgent health challenge facing humans today involves human-induced population bottlenecks that produce antibiotic-resistant bacteria. <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1451\"><strong>Antibiotics<\/strong> <\/a>are medicines prescribed to treat bacterial infections. The typical prescription includes enough medicine for ten days. People often feel better much sooner than ten days and sometimes decide to quit taking the medicine ahead of schedule. This is often a big mistake. The antibiotics have quickly killed off a large percentage of the bacteria\u2014enough to reduce the symptoms and make you feel much better. However, this has created a bacterial population bottleneck. There are usually a small number of bacteria that survive those early days. If you take the medicine as prescribed for the full ten days, it\u2019s quite likely that there will be no bacterial survivors. If you quit early, though, the survivors\u2014who were the members of the original population who were most resistant to the antibiotic\u2014will begin to reproduce again. Soon the infection will be back, possibly worse than before, and now all of the bacteria are resistant to the antibiotic that you had been prescribed.<\/p>\n<p class=\"import-Normal\">Other activities that have contributed to the rise of antibiotic-resistant bacteria include the use of antibacterial cleaning products and the inappropriate use of antibiotics as a preventative measure in livestock or to treat infections that are viral instead of bacterial (viruses do not respond to antibiotics). In 2017, the World Health Organization published a list of twelve antibiotic-resistant pathogens that are considered top priority targets for the development of new antibiotics (World Health Organization 2017).<\/p>\n<div class=\"textbox shaded\" style=\"background: var(--lightblue)\">\n<h2>Dig Deeper: The North American Elephant Seal: Thriving Bottleneck Populations That Still Face Genetic Defects<\/h2>\n<p>In 1892, the Northern Elephant Seal underwent a severe population bottleneck caused by commercial hunting, reducing the species to an estimated 20 individuals at the time. This drastic decline led to a substantial loss of genetic diversity\u2013a common consequence of extreme population bottlenecks (Hoelzel et al., 2024 &amp; Weber et al., 2000). While the population has since recovered to over 200,000 individuals, its genetic variability remains significantly low. Analyses of genetic markers, including allozymes, mitochondrial DNA, and microsatellites, consistently reflect this reduced diversity (Hoelzel et al., 2024). Comparative studies further underscore this loss by highlighting the higher genetic variation observed in the Southern Elephant Seal, which did not experience similar population constraints (2024).<\/p>\n<figure style=\"width: 386px\" class=\"wp-caption alignleft\"><img src=\"https:\/\/upload.wikimedia.org\/wikipedia\/commons\/thumb\/4\/48\/Elephant_seals_at_Ano_Nuevo_%2891577%29.jpg\/250px-Elephant_seals_at_Ano_Nuevo_%2891577%29.jpg\" alt=\"File:Elephant seals at Ano Nuevo (91577).jpg\" width=\"386\" height=\"295\" \/><figcaption class=\"wp-caption-text\"><span style=\"background-color: #ffff00\">Figure 5.24<\/span> A male northern elephant seal (Mirounga angustirostris) with two pups at Ano Nuevo State Park. Credit: Elephant seals at Ano Nuevo by Rhododendrites is under <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\" target=\"_blank\" rel=\"noopener\">Creative Commons Attribution-Share Alike 4.0<\/a>.<\/figcaption><\/figure>\n<p>In a 2024 study for Nature, Ecology, and Evolution, Hoelzel and colleagues sequenced 260 modern and 8 historical genomes of the northern elephant seal. This comparison revealed a decrease in average heterozygosity from 0.00142 before the bottleneck to 0.000176 in the contemporary population, confirming the decline in genetic variation (2024). Hoelzel\u2019s mitogenome tree further illustrates this loss, revealing only two significant lineages remaining post-bottleneck, with limited diversity within each. Among the issues of diversity, the population has shown an increased number of loss-of-function (LOF) alleles, suggesting that increased inbreeding has amplified the frequency of these detrimental alleles; this reduced genetic diversity negatively affects both male and female reproductive fitness. Females who practiced repetitive inbreeding had higher LOF alleles and subsequently weaned fewer pups per year over their lifetime, while male reproductive success was linked to specific LOF loci associated with sperm production (2024). Hoelzel uses the example of \u201cAlpha-Male M12\u201d\u2013known for low paternity success despite frequent copulations\u2013which was homozygous for non-functional versions of four out of five LOF loci related to sperm function (2024, p. 688). The species' mating system, characterized by extreme polygyny, further exacerbates the loss of genetic variation even with countless copulatory partners<\/p>\n<p>Prior research published in Current Biology presents an empirical genetic assessment of this population bottleneck, highlighting its long-term genetic consequences, particularly the loss of mitochondrial diversity (Weber et al., 2000). In this research, Weber and colleagues note that random lineage sampling during the bottleneck led to the persistence of specific genetic variants by chance rather than through natural selection (2000). This research emphasizes that the loss of diversity poses potential future genetic vulnerabilities for the seals, and that further studies are crucial for understanding the full scope of these impacts on the seals' overall fitness (2000). In 2024, the work led by Hoelzen and company provided the missing data that the previous study had left unanswered. Their previously explored findings indicate that, although the seals have recovered in numbers, their genetic resilience remains compromised, leaving the population more vulnerable to future environmental pressures, such as climate change or resource shortages (Hoelzel et al., 2024). Ultimately, while the population's size remains stable, the genetic consequences of the bottleneck indicate that past stochastic events continue to influence the seals' long-term fitness and adaptability.<\/p>\n<p>This research indicates that the historical bottleneck continues to affect the seals' health and fitness, despite the population's recovery. Limited genetic diversity and the persistence of harmful alleles due to inbreeding have continued to handicap the species' ability to thrive in environmental challenges such as climate change and resource fluctuations (2024). This emphasizes the importance of incorporating genetic factors into conservation strategies, as populations that have rebounded may still harbour long-term genetic weaknesses. Moreover, the elephant seal\u2019s history serves as a powerful example of how human actions \u2014such as overhunting \u2014 can have long-lasting impacts on biodiversity, reinforcing the importance of understanding human-environment interactions in ecological and conservation contexts.<\/p>\n<\/div>\n<h4 class=\"import-Normal\"><em>Founder Effects<\/em><\/h4>\n<p class=\"import-Normal\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1452\"><strong>Founder effects<\/strong><\/a> occur when members of a population leave the main or \u201cparent\u201d group and form a new population that no longer interbreeds with the other members of the original group. Similar to survivors of a population bottleneck, the newly founded population often has allele frequencies that are different from the original group. Alleles that may have been relatively rare in the parent population can end up being very common due to the founder effect. Likewise, recessive traits that were seldom seen in the parent population may be seen frequently in the descendants of the offshoot population.<\/p>\n<p class=\"import-Normal\">One striking example of the founder effect was first noted in the Dominican Republic in the 1970s. During a several-year period, eighteen children who had been born with female genitalia and raised as girls suddenly grew penises at puberty. This culture tended to value sons over daughters, so these transitions were generally celebrated. They labeled the condition <em><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1453\">guevedoces<\/a><\/strong><\/em>, which translates to \u201cpenis at twelve,\u201d due to the average age at which this occurred. Scientists were fascinated by the phenomenon.<\/p>\n<p class=\"import-Normal\">Genetic and hormonal studies revealed that the condition, scientifically termed <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1454\">5-alpha reductase deficiency<\/a>,<\/strong> is an autosomal recessive syndrome that manifests when a child having both X and Y sex chromosomes inherits two nonfunctional (mutated) copies of the <em>SRD5A2 <\/em>gene (Imperato-McGinley and Zhu 2002). These children develop testes internally, but the 5-alpha reductase 2 steroid, which is necessary for development of male genitals in babies, is not produced. In absence of this male hormone, the baby develops female-looking genitalia (in humans, \u201cfemale\u201d is the default infant body form, if the full set of the necessary male hormones are not produced). At puberty, however, a different set of male hormones are produced by other fully functional genes. These hormones complete the male genital development that did not happen in infancy. This condition became quite common in the Dominican Republic during the 1970s due to founder effect\u2014that is, the mutated <em>SRD5A2<\/em>\u00a0gene happened to be much more common among the Dominican Republic\u2019s founding population than in the parent populations. (The Dominican population derives from a mixture of Indigenous Americans [Taino] peoples, West Africans, and Western Europeans.) Five-alpha reductase syndrome has since been observed in other small, isolated populations around the world.<\/p>\n<p class=\"import-Normal\">Founder effect is closely linked to the concept of inbreeding, which in population genetics does not necessarily mean breeding with immediate family relatives. Instead, <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1455\"><strong>inbreeding<\/strong> <\/a> refers to the selection of mates exclusively from within a small, closed population\u2014that is, from a group with limited allelic variability. This can be observed in small, physically isolated populations but also can happen when cultural practices limit mates to a small group. As with the founder effect, inbreeding increases the risk of inheriting two copies of any nonfunctional (mutant) alleles.<\/p>\n<p class=\"import-Normal\">The Amish in the United States are a population that, due to their unique history and cultural practices, emerged from a small founding population and have tended to select mates from within their groups. The <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1456\">Old Order Amish<\/a><\/strong> population of Lancaster County, Pennsylvania, has approximately 50,000 current members, all of whom can trace their ancestry back to a group of approximately 80 individuals. This small founding population immigrated to the United States from Switzerland in the mid-1700s to escape religious persecution. Since the Amish keep to themselves and almost exclusively select mates from within their own communities, they have more recessive traits compared to their parent population.<\/p>\n<figure style=\"width: 441px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image11-1.jpg\" alt=\"One individual\u2019s hands with six fingers.\" width=\"441\" height=\"331\" \/><figcaption class=\"wp-caption-text\">Figure 5.11: A person displaying polydactyly. Credit: <a class=\"rId68\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:6_Finger.JPG\">6 Finger<\/a> by Wilhelmy is under a <a class=\"rId69\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">One of the genetic conditions that has been observed much more frequently in the Lancaster County Amish population is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1457\">Ellis-van Creveld syndrome<\/a><\/strong>, which is an autosomal recessive disorder characterized by short stature (dwarfism), polydactyly (the development of more than five digits [fingers or toes] on the hands or feet], abnormal tooth development, and heart defects (Figure 5.11). Among the general world population, Ellis-van Creveld syndrome is estimated to affect approximately 1 in 60,000 individuals; among the Old Order Amish of Lancaster County, the rate is estimated to be as high as 1 in every 200 births (D\u2019Asdia et al. 2013).<\/p>\n<p class=\"import-Normal\">One important insight that has come from the study of founder effects is that a limited gene pool carries a much higher risk for genetic diseases. Genetic diversity in a population greatly reduces these risks.<\/p>\n<h3 class=\"import-Normal\"><strong>Gene Flow<\/strong><\/h3>\n<p class=\"import-Normal\">The third force of evolution is traditionally called gene flow. As with genetic drift, this is a misnomer, because it refers to flowing alleles, not genes. (All members of the same species share the same genes; it is the alleles of those genes that may vary.) <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1459\"><strong>Gene <\/strong><strong>f<\/strong><strong>low<\/strong> <\/a> refers to the movement of alleles from one population to another. In most cases, gene flow can be considered synonymous with migration.<\/p>\n<p class=\"import-Normal\">Returning again to the example of our primordial cell population, let\u2019s imagine that, after the volcanic fissure opened up in the ocean floor, wiping out the majority of the parent population, two surviving populations developed in the waters on opposite sides of the fissure. Ultimately, the lava from the fissure cooled into a large island that continued to provide a physical barrier between the populations (Figure 5.12).<\/p>\n<figure style=\"width: 685px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image12-2.png\" alt=\"An illustration of gene flow.\" width=\"685\" height=\"342\" \/><figcaption class=\"wp-caption-text\">Figure 5.12: Smooth and predominantly ruffled amoeba-like populations separated by a volcanic eruption (at left) and an island (at right) with unidirectional gene flow moving from east to west with ocean currents. Credit: Population of amoeba-like cells separated by volcanic eruption original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Andrea J. Alveshere is a collective work under a <a class=\"rId74\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Includes <a class=\"rId75\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Amoeba_proteus_TK-UT.svg\">Amoeba Proteus TK-UT<\/a> by <a class=\"rId76\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Nefronus\">Tom\u00e1\u0161 Kebert<\/a> and <a class=\"rId77\" href=\"https:\/\/www.umimeto.org\/\">umimeto.org<\/a> (modified), <a class=\"rId78\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0<\/a>.]<\/figcaption><\/figure>\n<p class=\"import-Normal\">In the initial generations after the eruption, due to founder effect, isolation, and random inheritance (genetic drift), the population to the west of the islands contained a vast majority of the ruffled membrane alleles while the eastern population carried only the smooth alleles. Ocean currents in the area typically flowed from east to west, sometimes carrying cells (facilitating gene flow) from the eastern (smooth) population to the western (ruffled) population. Due to the ocean currents, it was almost impossible for any cells from the western population to be carried eastward. Thus, for inheritance purposes, the eastern (smooth) population remained isolated. In this case, the gene flow is unidirectional (going only in one direction) and unbalanced (only one population is receiving the new alleles).<\/p>\n<p class=\"import-Normal\">Among humans, gene flow is often described as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1460\"><strong>admixture<\/strong>.<\/a> In forensic cases, anthropologists and geneticists are often asked to estimate the ancestry of unidentified human remains to help determine whether they match any missing persons\u2019 reports. This is one of the most complicated tasks in these professions because, while \u201crace\u201d or \u201cancestry\u201d involves simple checkboxes on a missing person\u2019s form, among humans today there are no truly distinct genetic populations. All modern humans are members of the same fully breeding compatible species, and all human communities have experienced multiple episodes of gene flow (admixture), leading all humans today to be so genetically similar that we are all members of the same (and only surviving) human subspecies: <em>Homo sapiens sapiens.<\/em><\/p>\n<p class=\"import-Normal\">Gene flow between otherwise isolated nonhuman populations is often termed <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1461\">hybridization.<\/a>.<\/strong> One example of this involves the hybridization and spread of <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1462\"><strong>Scutellata<\/strong><strong> honey bees<\/strong><\/a> (a.k.a. \u201ckiller bees\u201d) in the Americas. All honey bees worldwide are classified as <em>Apis mellifera.<\/em> Due to distinct adaptations to various environments around the world, there are 28 different subspecies of <em>Apis mellifera<\/em>.<\/p>\n<p class=\"import-Normal\">During the 1950s, a Brazilian biologist named Warwick E. Kerr experimented with hybridizing African and European subspecies of honey bees to try to develop a strain that was better suited to tropical environments than the European honey bees that had long been kept by North American beekeepers. Dr. Kerr was careful to contain the reproductive queens and drones from the African subspecies, but in 1957, a visiting beekeeper accidentally released 26 queen bees of the Scutellata subspecies (<em>Apis mellifera scutellata<\/em>) from southern Africa into the Brazilian countryside. The Scutellata bees quickly interbred with local European honey bee populations. The hybridized bees exhibited a much more aggressively defensive behavior, fatally or near-fatally attacking many humans and livestock that ventured too close to their hives. The hybridized bees spread throughout South America and reached Mexico and California by 1985. By 1990, permanent colonies had been established in Texas, and by 1997, 90% of trapped bee swarms around Tucson, Arizona, were found to be Scutellata hybrids (Sanford 2006).<\/p>\n<p class=\"import-Normal\">Another example involves the introduction of the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1463\"><strong>Harlequin ladybeetle<\/strong><\/a>, <em>Harmonia axyridis<\/em>, native to East Asia, to other parts of the world as a \u201cnatural\u201d form of pest control. Harlequin ladybeetles are natural predators of some of the aphids and other crop-pest insects. First introduced to North America in 1916, the \u201cbiocontrol\u201d strains of Harlequin ladybeetles were considered to be quite successful in reducing crop pests and saving farmers substantial amounts of money. After many decades of successful use in North America, biocontrol strains of Harlequin ladybeetles were also developed in Europe and South America in the 1980s.<\/p>\n<p class=\"import-Normal\">Over the seven decades of biocontrol use, the Harlequin ladybeetle had never shown any potential for development of wild colonies outside of its native habitat in China and Japan. New generations of beetles always had to be reared in the lab. That all changed in 1988, when a wild colony took root near New Orleans, Louisiana. Either through admixture with a native ladybeetle strain, or due to a spontaneous mutation, a new allele was clearly introduced into this population that suddenly enabled them to survive and reproduce in a wide range of environments. This population spread rapidly across the Americas and had reached Africa by 2004.<\/p>\n<p class=\"import-Normal\">In Europe, the invasive, North American strain of Harlequin ladybeetle admixed with the European strain (Figure 5.13), causing a population explosion (Lombaert et al. 2010). Even strains specifically developed to be flightless (to curtail the spreading) produced flighted offspring after admixture with members of the North American population (Facon et al. 2011). The fast-spreading, invasive strain has quickly become a disaster, out-competing native ladybeetle populations (some to the point of extinction), causing home infestations, decimating fruit crops, and contaminating many batches of wine with their bitter flavor after being inadvertently harvested with the grapes (Pickering et al. 2004).<\/p>\n<figure style=\"width: 583px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image13-2.png\" alt=\"One gray ladybug is migrating to the group of white ladybugs.\" width=\"583\" height=\"219\" \/><figcaption class=\"wp-caption-text\">Figure 5.13: Gene flow between two populations of ladybeetles (ladybugs). Credit: <a class=\"rId80\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Ladybug Gene Flow (Figure 4.14)<\/a> original to <a class=\"rId81\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a class=\"rId82\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Natural Selection<\/strong><\/h3>\n<p class=\"import-Normal\">The final force of evolution is natural selection. This is the evolutionary process that Charles Darwin first brought to light, and it is what the general public typically evokes when considering the process of evolution. <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1465\"><strong>Natural <\/strong><strong>s<\/strong><strong>election<\/strong><\/a> occurs when certain phenotypes confer an advantage or disadvantage in survival and\/or reproductive success. The alleles associated with those phenotypes will change in frequency over time due to this selective pressure. It\u2019s also important to note that the advantageous allele may change over time (with environmental changes) and that an allele that had previously been benign may become advantageous or detrimental. Of course, dominant, recessive, and codominant traits will be selected upon a bit differently from one another. Because natural selection acts on phenotypes rather than the alleles themselves, deleterious (disadvantageous) alleles can be retained by heterozygotes without any negative effects.<\/p>\n<p class=\"import-Normal\">In the case of our primordial ocean cells, up until now, the texture of their cell membranes has been benign. The frequencies of smooth to ruffled alleles, and smooth to ruffled phenotypes, has changed over time, due to genetic drift and gene flow. Let\u2019s now imagine that the Earth\u2019s climate has cooled to a point that the waters frequently become too cold for survival of the tiny bacteria that are the dietary staples of our smooth and ruffled cell populations. The way amoeba-like cells \u201ceat\u201d is to stretch out the cell membrane, almost like an arm, to encapsulate, then ingest, the tiny bacteria. When the temperatures plummet, the tiny bacteria populations plummet with them. Larger bacteria, however, are better able to withstand the temperature change.<\/p>\n<p class=\"import-Normal\">The smooth cells were well-adapted to ingesting tiny bacteria but poorly suited to encapsulating the larger bacteria. The cells with the ruffled membranes, however, are easily able to extend their ruffles to encapsulate the larger bacteria. They also find themselves able to stretch their entire membrane to a much larger size than their smooth-surfaced neighbors, allowing them to ingest more bacteria at a given time and to go for longer periods between feedings (Figure 5.14).<\/p>\n<figure style=\"width: 528px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image14-2.png\" alt=\"Smooth and ruffled cells feeding on large and small bacteria.\" width=\"528\" height=\"307\" \/><figcaption class=\"wp-caption-text\">Figure 5.14: Smooth and ruffled cells feeding. Credit: Smooth and ruffled cells feeding original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Andrea J. Alveshere is a collective work under a <a class=\"rId84\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>. [Includes <a class=\"rId85\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Cladograma_dos_Dom%C3%ADnios_e_Reinos.png\">Cladograma dos Dominios e Reinos<\/a> by <a class=\"rId86\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:MarceloTeles\">MarceloTeles<\/a> (modified), <a class=\"rId87\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0<\/a><a class=\"rId88\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">; <\/a><a class=\"rId89\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Amoeba_proteus_TK-UT.svg\">Amoeba Proteus TK-UT<\/a> by <a class=\"rId90\" href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Nefronus\">Tom\u00e1\u0161 Kebert<\/a> and <a class=\"rId91\" href=\"https:\/\/www.umimeto.org\/\">umimeto.org<\/a> (modified), <a class=\"rId92\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/deed.en\">CC BY-SA 4.0<\/a>.]<\/figcaption><\/figure>\n<p class=\"import-Normal\">The smooth and ruffled traits, which had previously offered no advantage or disadvantage while food was plentiful, now are subject to natural selection. During the cold snaps, at least, the ruffled cells have a definite advantage. We can imagine that the western population that has mostly ruffled alleles will continue to do well, while the eastern population is at risk of dying out if the smaller bacteria remain scarce and no ruffled alleles are introduced.<\/p>\n<p class=\"import-Normal\">A classic example of natural selection involves the study of an insect called the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1466\">peppered moth<\/a><\/strong> (<em>Biston betularia<\/em>) in England during the Industrial Revolution in the 1800s. Prior to the Industrial Revolution, the peppered moth population was predominantly light in color, with dark (pepper-like) speckles on the wings. The \u201cpeppered\u201d coloration was very similar to the appearance of the bark and lichens that grew on the local trees (Figure 5.15). This helped to camouflage the moths as they rested on a tree, making it harder for moth-eating birds to find and snack on them. There was another phenotype that popped up occasionally in the population. These individuals were heterozygotes that carried an overactive, dominant pigment allele, producing a solid black coloration. As you can imagine, the black moths were much easier for birds to spot, making this phenotype a real disadvantage.<\/p>\n<p class=\"import-Normal\">The situation changed, however, as the Industrial Revolution took off. Large factories began spewing vast amounts of coal smoke into the air, blanketing the countryside, including the lichens and trees, in black soot. Suddenly, it was the light-colored moths that were easy for birds to spot and the black moths that held the advantage. The frequency of the dark pigment allele rose dramatically. By 1895, the black moth phenotype accounted for 98% of observed moths (Grant 1999).<\/p>\n<figure style=\"width: 476px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image15-2.png\" alt=\"An illustration of natural selection.\" width=\"476\" height=\"531\" \/><figcaption class=\"wp-caption-text\">Figure 5.15: Dark and light peppered moth variants and their relative camouflage abilities on clean (top) and sooty (bottom) trees. Credit: <a class=\"rId94\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Peppered_moths_c2.jpg\">Peppered moths c2<\/a> by Khaydock is under a <a class=\"rId95\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Thanks to new environmental regulations in the 1960s, the air pollution in England began to taper off. As the soot levels decreased, returning the trees to their former, lighter color, this provided the perfect opportunity to study how the peppered moth population would respond. Repeated follow-up studies documented the gradual rise in the frequency of the lighter-colored phenotype. By 2003, the maximum frequency of the dark phenotype was 50% and in most parts of England had decreased to less than 10% (Cook 2003).<\/p>\n<h4 class=\"import-Normal\"><em>Directional, Balancing\/Stabilizing, and Disruptive\/Diversifying Selection<\/em><\/h4>\n<p class=\"import-Normal\">Natural selection can be classified as directional, balancing\/stabilizing, or disruptive\/diversifying, depending on how the pressure is applied to the population (Figure 5.16).<\/p>\n<figure style=\"width: 465px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image16-2.png\" alt=\"Three types of selection; balancing, directional and disruptive\/diversifying\" width=\"465\" height=\"574\" \/><figcaption class=\"wp-caption-text\">Figure 5.16: Lines depict the affects of (a) Balancing\/Stabilizing, (b) Directional, and (c) Disruptive\/Diversifying selection on populations. <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 class=\"rId97\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Figure_19_03_01.png\">Biology (ID: 185cbf87-c72e-48f5-b51e-f14f21b5eabd@9.17)<\/a> by <a class=\"rId98\" href=\"https:\/\/cnx.org\/\">CNX OpenStax<\/a> is used under a <a class=\"rId99\" href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/deed.en\">CC BY 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Both of the above examples of natural selection involve <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1467\">directional selection<\/a><\/strong>: the environmental pressures favor one phenotype over the other and cause the frequencies of the associated advantageous alleles (ruffled membranes, dark pigment) to gradually increase. In the case of the peppered moths, the direction shifted three times: first, it was selecting for lighter pigment; then, with the increase in pollution, the pressure switched to selection for darker pigment; finally, with reduction of the pollution, the selection pressure shifted back again to favoring light-colored moths.<\/p>\n<p class=\"import-Normal\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1468\"><strong>Balancing selection<\/strong><\/a> (a.k.a. stabilizing selection) occurs when selection works against the extremes of a trait and favors the intermediate phenotype. For example, humans maintain an average birth weight that balances the need for babies to be small enough not to cause complications during pregnancy and childbirth but big enough to maintain a safe body temperature after they are born. Another example of balancing selection is found in the genetic disorder called sickle cell anemia (see \u201cSpecial Topic: Sickle Cell Anemia\u201d).<\/p>\n<p class=\"import-Normal\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1469\"><strong>Disruptive selection<\/strong><\/a> (a.k.a. diversifying selection), the opposite of balancing selection, occurs when both extremes of a trait are advantageous. Since individuals with traits in the mid-range are selected against, disruptive selection can eventually lead to the population evolving into two separate species. Darwin believed that the many species of finches (small birds) found in the remote Galapagos Islands provided a clear example of disruptive selection leading to speciation. He observed that seed-eating finches either had large beaks, capable of eating very large seeds, or small beaks, capable of retrieving tiny seeds. The islands did not have many plants that produced medium-size seeds. Thus, birds with medium-size beaks would have trouble eating the very large seeds and would also have been inefficient at picking up the tiny seeds. Over time, Darwin surmised, this pressure against mid-size beaks may have led the population to divide into two separate species.<\/p>\n<h4 class=\"import-Normal\"><em>Sexual Selection<\/em><\/h4>\n<p class=\"import-Normal\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1470\"><strong>Sexual <\/strong><strong>s<\/strong><strong>election<\/strong><\/a> is an aspect of natural selection in which the selective pressure specifically affects reproductive success (the ability to successfully breed and raise offspring) rather than survival. Sexual selection favors traits that will attract a mate. Sometimes these sexually appealing traits even carry greater risks in terms of survival.<\/p>\n<figure style=\"width: 354px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image17-2.png\" alt=\"A fox chases a peacock fleeing; a peacock displays his feathers to a peahen.\" width=\"354\" height=\"413\" \/><figcaption class=\"wp-caption-text\">Figure 5.17: Showy peacock tail disadvantages (becoming easier prey) and advantages (impressing peahens). Credit: <a class=\"rId101\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Peacock tail advantage and disadvantages (Figure 4.18)<\/a> original to <a class=\"rId102\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a class=\"rId103\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.License.<\/figcaption><\/figure>\n<p class=\"import-Normal\">A classic example of sexual selection involves the brightly colored feathers of the peacock. The <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1471\"><strong>peacock<\/strong><\/a> is the male sex of the peafowl genera <em>Pavo<\/em>\u00a0and\u00a0<em>Afropavo. <\/em>During mating season, peacocks will fan their colorful tails wide and strut in front of the peahens in a grand display. The peahens will carefully observe these displays and will elect to mate with the male that they find the most appealing. Many studies have found that peahens prefer the males with the fullest, most colorful tails. While these large, showy tails provide a reproductive advantage, they can be a real burden in terms of escaping predators. The bright colors and patterns as well as the large size of the peacock tail make it difficult to hide. Once predators spot them, peacocks also struggle to fly away, with the heavy tail trailing behind and weighing them down (Figure 5.17). Some researchers have argued that the increased risk is part of the appeal for the peahens: only an especially strong, alert, and healthy peacock would be able to avoid predators while sporting such a spectacular tail.<\/p>\n<\/div>\n<p>It\u2019s important to keep in mind that sexual selection relies on the trait being present throughout mating years. Reflecting on the NF1 genetic disorder (see \u201cSpecial Topic: Neurofibromatosis Type 1 [NF1]\u201d), given how disfiguring the symptoms can become, some might find it surprising that half of the babies born with NF1 inherited it from a parent. Given that the disorder is autosomal dominant and fully penetrant (meaning it has no unaffected carriers), it may seem surprising that sexual selection doesn\u2019t exert more pressure against the mutated alleles. One important factor is that, while the neurofibromas typically begin to appear during puberty, they usually emerge only a few at a time and may grow very slowly. Many NF1 patients don\u2019t experience the more severe or disfiguring symptoms until later in life, long after they have started families of their own.<\/p>\n<p class=\"import-Normal\">Some researchers prefer to classify sexual selection separately, as a fifth force of evolution. The traits that underpin mate selection are entirely natural, of course. Research has shown that subtle traits, such as the type of pheromones (hormonal odors related to immune system alleles) someone emits and how those are perceived by the immune system genotype of the \u201csniffer,\u201d may play crucial and subconscious roles in whether we find someone attractive or not (Chaix, Cao, and Donnelly 2008).<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Neurofibromatosis Type 1 (NF1)<\/h2>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1476\">Neurofibromatosis Type 1<\/a><\/strong>, also known as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1477\"><strong>NF1<\/strong><\/a>, is a genetic disorder that illustrates how a mutation in a single gene can affect multiple systems in the body. Surprisingly common, more people have NF1 than cystic fibrosis and muscular dystrophy combined. Even more surprising, given how common it is, is how few people have heard of it. One in every 3,000 babies is born with NF1, and this holds true for all populations worldwide (Riccardi 1992). This means that, for every 3,000 people in your community, there is likely at least one person living with this disorder. NF1 is an <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1478\">autosomal dominant<\/a> <\/strong>condition, which means that everyone born with a mutation in the gene, whether inherited or spontaneous, has a 50\/50 chance of passing it on to each of their own children.<\/p>\n<p class=\"import-Normal\">The NF1 disorder results from mutation of the <em>NF1<\/em> gene on Chromosome 17. Almost any mutation that affects the sequence of the gene\u2019s protein product, neurofibromin, will cause the disorder. Studies of individuals with NF1 have identified over 3,000 different mutations of all kinds (including point mutations, small and large indels, and translocations). The <em>NF1 <\/em>gene is one of the largest known genes, containing at least 60 <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_724\">exons<\/a><\/strong> (protein-encoding sequences) in a span of about 300,000 nucleotides.<\/p>\n<p class=\"import-Normal\">We know that neurofibromin plays an important role in preventing tumor growth because one of the most common symptoms of the NF1 disorder is the growth of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1479\">benign<\/a> <\/strong>(noncancerous) tumors, called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1486\">neurofibromas<\/a><\/strong>. Neurofibromas sprout from nerve sheaths\u2014the tissues that encase our nerves\u2014throughout the body, usually beginning around puberty. There is no way to predict where the tumors will occur, or when or how quickly they will grow, although only about 15% turn <strong>malignant<\/strong> (cancerous). The two types of neurofibromas that are typically most visible are <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1480\"><strong>cutaneous neurofibromas<\/strong><\/a>, which are spherical bumps on, or just under, the surface of the skin (Figure 5.18), and <strong>plexiform neurofibromas<\/strong><em>, <\/em>growths involving whole branches of nerves, often giving the appearance that the surface of the skin is \u201cmelting\u201d (Figure 5.19).<\/p>\n<figure id=\"attachment_131-3\" aria-describedby=\"caption-attachment-131-3\" style=\"width: 510px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-129\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/4.18.jpg\" alt=\"A woman has dozens of round, skin-colored tumors visible on her face, neck, and hand.\" width=\"510\" height=\"340\" \/><figcaption id=\"caption-attachment-131-3\" class=\"wp-caption-text\">Figure 5.18: A woman with many cutaneous neurofibromas, a common symptom of Neurofibromatosis Type 1. Credit: <a class=\"rId105\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Woman with cutaneous neurofibromas (symptom of NF1)<\/a> by <a class=\"rId106\" href=\"https:\/\/positiveexposure.org\/about-the-program-2\/rick-guidotti\/\">Rick Guidotti of Positive Exposure<\/a> is used with permission and is available here under a <a class=\"rId107\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<figure id=\"attachment_131-4\" aria-describedby=\"caption-attachment-131-4\" style=\"width: 1900px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-130 size-full\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/4.19.jpg\" alt=\"An adult with large plexiform neurofibromas covering his face, none are on the child.\" width=\"1900\" height=\"700\" \/><figcaption id=\"caption-attachment-131-4\" class=\"wp-caption-text\">Figure 5.19: Photo on the left is of a man with large plexiform neurofibroma, another symptom of Neurofibromatosis Type 1. Photo on the right is a childhood photo of the same man, illustrating the progressive nature of the NF1 disorder. Credit: <a class=\"rId110\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Man with plexiform neurofibroma (symptom of NF1)<\/a> from Ashok Shrestha is used by permission and available here under a <a class=\"rId111\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. <a class=\"rId112\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Childhood photo of the same man with NF1 disorder<\/a> from Ashok Shrestha is used by permission and available here under a <a class=\"rId113\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Unfortunately, there is currently no cure for NF1. Surgical removal of neurofibromas risks paralysis, due to the high potential for nerve damage, and often results in the tumors growing back even more vigorously. This means that patients are often forced to live with disfiguring and often painful neurofibromas. People who are not familiar with NF1 often mistake neurofibromas for something contagious. This makes it especially hard for people living with NF1 to get jobs working with the public or even to enjoy spending time away from home. Raising public awareness about NF1 and its symptoms can be a great help in improving the quality of life for people living with this condition.<\/p>\n<figure style=\"width: 311px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image21-2.png\" alt=\"A child with darker oval birthmarks scattered across his torso and arms.\" width=\"311\" height=\"415\" \/><figcaption class=\"wp-caption-text\">Figure 5.20: Image of a child with caf\u00e9-au-lait macules (birthmarks) typical of the earliest symptoms of NF1. Credit: <a class=\"rId115\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Child with caf\u00e9-au-lait macules (birthmarks) typical of the earliest symptoms of NF1<\/a> by Andrea J. Alveshere is under a <a class=\"rId116\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">One of the first symptoms of NF1 in a small child is usually the appearance of <strong>caf\u00e9-au-lait spots<\/strong>, or <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1490\"><strong>CALS<\/strong><\/a>, which are flat, brown birthmark-like spots on the skin (Figure 5.20). CALS are often light brown, similar to the color of coffee with cream, which is the reason for the name, although the shade of the pigment depends on a person\u2019s overall complexion. Some babies are born with CALS, but for others the spots appear within the first few years of life. Having six or more CALS larger than five millimeters (mm) across is a strong indicator that a child may have NF1.<\/p>\n<p class=\"import-Normal\">Other common symptoms include the following: gliomas (tumors) of the optic nerve, which can cause vision loss; thinning of bones and failure to heal if they break (often requiring amputation); low muscle tone (poor muscle development, often delaying milestones such as sitting up, crawling, and walking); hearing loss, due to neurofibromas on auditory nerves; and learning disabilities, especially those involving spatial reasoning. Approximately 50% of people with NF1 have some type of speech and\/or learning disability and often benefit greatly from early intervention services. Generalized developmental disability, however, is not common with NF1, so most people with NF1 live independently as adults. Many people with NF1 live full and successful lives, as long as their symptoms can be managed.<\/p>\n<p class=\"import-Normal\">Based on the wide variety of symptoms, it\u2019s clear that the neurofibromin protein plays important roles in many biochemical pathways. While everyone who has NF1 will exhibit some symptoms during their lifetime, there is a great deal of variation in the types and severity of symptoms, even between individuals from the same family who share the exact same NF1 mutation. It seems crazy that a gene with so many important functions would be so susceptible to mutation. Part of this undoubtedly has to do with its massive size\u2014a gene with 300,000 nucleotides has ten times more nucleotides available for mutation than does a gene of 30,000 bases. This also suggests that the mutability of this gene might provide some benefits, which is a possibility that we will revisit later in this chapter.<\/p>\n<\/div>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Sickle Cell Anemia<\/h2>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1492\">Sickle cell anemia<\/a><\/strong> is an autosomal recessive genetic disorder that affects millions of people worldwide. It is most common in Africa, countries around the Mediterranean Sea, and eastward as far as India. Populations in the Americas that have high percentages of ancestors from these regions also have high rates of sickle cell anemia. In the United States, it\u2019s estimated that 72,000 people live with the disease, with one in approximately 1,200 Hispanic-American babies and one in every 500 African-American babies inheriting the condition (World Health Organization 1996).<\/p>\n<figure style=\"width: 344px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image22-2.png\" alt=\"Round and sickle cells.\" width=\"344\" height=\"258\" \/><figcaption class=\"wp-caption-text\">Figure 5.21: Sickle cell anemia. Arrows indicate (a) sickled and (b) normal red blood cells. Credit: <a class=\"rId118\" href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Sickle-cell_smear_2015-09-10.jpg\">Sickle-cell smear 2015-09-10<\/a> by Paulo Henrique Orlandi Mourao has been modified (contrast modified and labels added) and is under a <a class=\"rId119\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Sickle cell anemia affects the hemoglobin protein in red blood cells. Normal red blood cells are somewhat doughnut-shaped\u2014round with a depression on both sides of the middle. They carry oxygen around the bloodstream to cells throughout the body. Red blood cells produced by the mutated form of the gene take on a stiff, sickle-like crescent shape when stressed by low oxygen or dehydration (Figure 5.21). Because of their elongated shape and the fact that they are stiff rather than flexible, they tend to form clumps in the blood vessels, inhibiting blood flow to adjacent areas of the body. This causes episodes of extreme pain and can cause serious problems in the oxygen-deprived tissues. The sickle cells also break down much more quickly than normal cells, often lasting only 20 days rather than the 120 days of normal cells. This causes an overall shortage of blood cells in the sickle cell patient, resulting in low iron (anemia) and problems associated with it such as extreme fatigue, shortness of breath, and hindrances to children\u2019s growth and development.<\/p>\n<p class=\"import-Normal\">The devastating effects of sickle cell anemia made its high frequency a pressing mystery. Why would an allele that is so deleterious in its homozygous form be maintained in a population at levels as high as the one in twelve African Americans estimated to carry at least one copy of the allele? The answer turned out to be one of the most interesting cases of balancing selection in the history of genetic study.<\/p>\n<p class=\"import-Normal\">While looking for an explanation, scientists noticed that the countries with high rates of sickle cell disease also shared a high risk for another disease called <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1493\"><strong>malaria<\/strong><\/a>, which is caused by infection of the blood by a <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1494\"><strong><em>Plasmodium<\/em><\/strong><\/a> parasite. These parasites are carried by mosquitoes and enter the human bloodstream via a mosquito bite. Once infected, the person will experience flu-like symptoms that, if untreated, can often lead to death. Researchers discovered that many people living in these regions seemed to have a natural resistance to malaria. Further study revealed that people who carry the sickle cell allele are far less likely to experience a severe case of malaria. This would not be enough of a benefit to make the allele advantageous for the sickle cell homozygotes, who face shortened life spans due to sickle cell anemia. The real benefit of the sickle cell allele goes to the heterozygotes.<\/p>\n<p class=\"import-Normal\">People who are heterozygous for sickle cell carry one normal allele, which produces the normal, round, red blood cells, and one sickle cell allele, which produces the sickle-shaped red blood cells. Thus, they have both the sickle and round blood cell types in their bloodstream. They produce enough of the round red blood cells to avoid the symptoms of sickle cell anemia, but they have enough sickle cells to provide protection from malaria.<\/p>\n<p class=\"import-Normal\">When the <em>Plasmodium <\/em>parasites infect an individual, they begin to multiply in the liver, but then must infect the red blood cells to complete their reproductive cycle. When the parasites enter sickle-type cells, the cells respond by taking on the sickle shape. This prevents the parasite from circulating through the bloodstream and completing its life cycle, greatly inhibiting the severity of the infection in the sickle cell heterozygotes compared to non\u2013-sickle cell homozygotes. See Chapter 14 for more discussion of sickle cell anemia.<\/p>\n<\/div>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: The Real Primordial Cells\u2014<em>Dictyostelium Discoideum<\/em><\/h2>\n<p class=\"import-Normal\">The amoeba-like primordial cells that were used as recurring examples throughout this chapter are inspired by actual research that is truly fascinating. In 2015, Gareth Bloomfield and colleagues reported on their genomic study of the social amoeba <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1496\"><strong><em>Dictyostelium discoideum<\/em><\/strong> <\/a>(a.k.a. \u201cslime molds,\u201d although technically they are amoebae, not molds). Strains of these amoebae have been grown in research laboratories for many decades and are useful in studying the mechanisms that amoeboid single-celled organisms use to ingest food and liquid. For simplification of our examples in this chapter, our amoeba-like cells remained ocean dwellers. Wild <em>Dictyostelium discoideum<\/em>, however, live in soil and feed on soil bacteria by growing ruffles in their membranes that reach out to encapsulate the bacterial cell. Laboratory strains, however, are typically raised on liquid media (agar) in Petri dishes, which is not suitable for the wild-type amoebae. It was widely known that the laboratory strains must have developed mutations in one or more genes to allow them to ingest the larger nutrient particles in the agar and larger volumes of liquid, but the genes involved were not known.<\/p>\n<p class=\"import-Normal\">Bloomfield and colleagues performed genomic testing on both the wild and the laboratory strains of <em>Dictyostelium discoideum. <\/em>Their discovery was astounding: every one of the laboratory strains carried a mutation in the <em>NF1 <\/em>gene, the very same gene associated with Neurofibromatosis Type 1 (NF1) in humans. The antiquity of this massive, easily mutated gene is incredible. It originated in an ancestor common to both humans and these amoebae, and it has been retained in both lineages ever since. As seen in <em>Dictyostelium discoideum<\/em>, breaking the gene can be advantageous. Without a functioning copy of the neurofibromin protein, the cell membrane is able to form much-larger feeding structures, allowing the <em>NF1 <\/em>mutants to ingest larger particles and larger volumes of liquid. For these amoebae, this may provide dietary flexibility that functions somewhat like an insurance policy for times when the food supply is limited.<\/p>\n<p class=\"import-Normal\"><em>Dictyostelium discoideum <\/em>are also interesting in that they typically reproduce asexually, but under certain conditions, one cell will convert into a \u201cgiant\u201d cell, which encapsulates surrounding cells, transforming into one of three sexes. This cell will undergo meiosis, producing gametes that must combine with one of the other two sexes to produce viable offspring. This ability for sexual reproduction may be what allows <em>Dictyostelium discoideum<\/em> to benefit from the advantages of <em>NF1<\/em> mutation, while also being able to restore the wild type <em>NF1<\/em> gene in future generations.<\/p>\n<p class=\"import-Normal\">What does this mean for humans living with NF1? Well, understanding the role of the neurofibromin protein in the membranes of simple organisms like <em>Dictyostelium discoideum<\/em> may help us to better understand how it functions and malfunctions in the sheaths of human neurons. It\u2019s also possible that the mutability of the NF1 gene confers certain advantages to humans as well. Alleles of the NF1 gene have been found to reduce one\u2019s risk for alcoholism (Repunte-Canonigo Vez et al. 2015), opiate addiction (Sanna et al. 2002), Type 2 diabetes (Martins et al. 2016), and hypomusicality (a lower-than-average musical aptitude; Cota et al. 2018). This research is ongoing and will be exciting to follow in the coming years.<\/p>\n<\/div>\n<h2 class=\"__UNKNOWN__\">Studying Evolution in Action<\/h2>\n<div class=\"__UNKNOWN__\">\n<h3 class=\"import-Normal\"><strong>The Hardy-Weinberg Equilibrium <\/strong><\/h3>\n<p class=\"import-Normal\">This chapter has introduced you to the forces of evolution, the mechanisms by which evolution occurs. How do we detect and study evolution, though, in real time, as it happens? One tool we use is the <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1497\"><strong>Hardy-<\/strong><strong>Weinberg<\/strong><strong> Equilibrium<\/strong><\/a>: a mathematical formula that allows estimation of the number and distribution of dominant and recessive alleles in a population. This aids in determining whether allele frequencies are changing and, if so, how quickly over time, and in favor of which allele? It\u2019s important to note that the Hardy-Weinberg formula only gives us an estimate based on the data for a snapshot in time. We will have to calculate it again later, after various intervals, to determine if our population is evolving and in what way the allele frequencies are changing.<\/p>\n<h3 class=\"import-Normal\">Calculating the Hardy-Weinberg Equilibrium<\/h3>\n<p class=\"import-Normal\">In the Hardy-Weinberg formula, <em>p <\/em>represents the frequency of the dominant allele, and <em>q<\/em> represents the frequency of the recessive allele. Remember, an allele\u2019s frequency is the proportion, or percentage, of that allele in the population. For the purposes of Hardy-Weinberg, we give the allele percentages as decimal numbers (e.g., 42% = 0.42), with the entire population (100% of alleles) equaling 1. If we can figure out the frequency of one of the alleles in the population, then it is simple to calculate the other. Simply subtract the known frequency from 1 (the entire population): 1<em> \u2013 p = q<\/em> and 1<em> \u2013 q = p<\/em>.<\/p>\n<p class=\"import-Normal\">The Hardy-Weinberg formula is <em>p<\/em><sup><em>2<\/em><\/sup><em> + 2pq + q<\/em><sup><em>2<\/em><\/sup>, where:<\/p>\n<p class=\"import-Normal\" style=\"padding-left: 40px\"><em>p<\/em><sup><em>2<\/em><\/sup> represents the frequency of the homozygous dominant genotype;<\/p>\n<p class=\"import-Normal\" style=\"padding-left: 40px\"><em>2pq<\/em> represents the frequency of the heterozygous genotype; and<\/p>\n<p class=\"import-Normal\" style=\"padding-left: 40px\"><em>q<\/em><sup><em>2<\/em><\/sup> represents the frequency of the homozygous recessive genotype.<\/p>\n<p class=\"import-Normal\">It is often easiest to determine <em>q<\/em><sup><em>2<\/em><\/sup> first, simply by counting the number of individuals with the unique, homozygous recessive phenotype (then dividing by the total individuals in the population to arrive at the \u201cfrequency\u201d). Once we have this number, we simply need to calculate the square root of the homozygous recessive phenotype frequency. That gives us <em>q.<\/em> Remember, 1 <em>\u2013<\/em> <em>q <\/em>equals <em>p<\/em>, so now we have the frequencies for both alleles in the population. If we needed to figure out the frequencies of heterozygotes and homozygous dominant genotypes, we\u2019d just need to plug the <em>p<\/em> and <em>q<\/em> frequencies back into the <em>p<\/em><sup><em>2<\/em><\/sup> and 2<em>pq<\/em> formulas.<\/p>\n<figure style=\"width: 329px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image25-1.png\" alt=\"A circle with seven grey and three white ladybugs.\" width=\"329\" height=\"347\" \/><figcaption class=\"wp-caption-text\">Figure 5.24: Ladybug population with a mixture of dark (red) and light (orange) individuals. Credit: <a class=\"rId129\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Ladybug mix (Figure 4.21)<\/a> original to <a class=\"rId130\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a class=\"rId131\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Let\u2019s imagine we have a population of ladybeetles that carries two alleles: a dominant allele that produces red ladybeetles and a recessive allele that produces orange ladybeetles. Since red is dominant, we\u2019ll use <em>R <\/em>to represent the red allele, and <em>r <\/em>to represent the orange allele. Our population has ten beetles, and seven are red and three are orange (Figure 5.24). Let\u2019s calculate the number of genotypes and alleles in this population.<\/p>\n<p class=\"import-Normal\">Of ten total beetles, we have three orange beetles3\/10 = .30 (30%) frequency\u2014and we know they are homozygous recessive (<em>rr<\/em>). So:<\/p>\n<p class=\"import-Normal\" style=\"text-indent: 36pt\"><em>rr = .3; <\/em>therefore, <em>r = <\/em>\u221a.3 = .5477<\/p>\n<p class=\"import-Normal\" style=\"text-indent: 36pt\"><em>R<\/em> = 1 <em>\u2013<\/em> .5477 = .4523<\/p>\n<p class=\"import-Normal\" style=\"text-indent: 36pt\">Using the Hardy-Weinberg formula:<\/p>\n<p class=\"import-Normal\" style=\"text-indent: 36pt\">1=.4523<sup>2 <\/sup>+ 2 x .4523 x .5477 +.5477<sup>2 <\/sup>= .20 + .50 + .30 = 1<\/p>\n<p class=\"import-Normal\" style=\"text-indent: 36pt\">Thus, the genotype breakdown is 20% <em>RR, <\/em>50%<em> Rr, <\/em>and 30%<em> rr <\/em><\/p>\n<p class=\"import-Normal\" style=\"text-indent: 36pt\">(2 red homozygotes, 5 red heterozygotes, and 3 orange homozygotes).<\/p>\n<p class=\"import-Normal\">Since we have 10 individuals, we know we have 20 total alleles: 4 red from the <em>RR<\/em> group, 5 red and 5 orange from the <em>Rr<\/em> group, and 6 orange from the <em>rr<\/em> group, for a grand total of 9 red and 11 orange (45% red and 55% orange, just like we estimated in the 1 \u2013 <em>q <\/em>step).<\/p>\n<p class=\"import-Normal\">Reminder: The Hardy-Weinberg formula only gives us an estimate for a snapshot in time. We will have to calculate it again later, after various intervals, to determine if our population is evolving and in what way the allele frequencies are changing.<\/p>\n<h3 class=\"import-Normal\"><strong>Interpreting Evolutionary Change: Nonra<\/strong><strong>ndom Mating <\/strong><\/h3>\n<p class=\"import-Normal\">Once we have detected change occurring in a population, we need to consider which evolutionary processes might be the cause of the change. It is important to watch for nonrandom mating patterns, to see if they can be included or excluded as possible sources of variation in allele frequencies.<\/p>\n<p class=\"import-Normal\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1498\"><strong>Nonrandom <\/strong><strong>m<\/strong><strong>ating<\/strong><\/a> (also known as assortative mating) occurs when mate choice within a population follows a nonrandom pattern.<\/p>\n<p class=\"import-Normal\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1499\"><strong>Positive assortative mating<\/strong><\/a> patterns result from a tendency for individuals to mate with others who share similar phenotypes. This often happens based on body size. Taking as an example dog breeds, it is easier for two Chihuahuas to mate and have healthy offspring than it is for a Chihuahua and a St. Bernard to do so. This is especially true if the Chihuahua is the female and would have to give birth to giant St. Bernard pups.<\/p>\n<p class=\"import-Normal\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1500\"><strong>Negative assortative mating<\/strong><\/a> patterns occur when individuals tend to select mates with qualities different from their own. This is what is at work when humans choose partners whose pheromones indicate that they have different and complementary immune alleles, providing potential offspring with a better chance at a stronger immune system.<\/p>\n<p class=\"import-Normal\">Among domestic animals, such as pets and livestock, assortative mating is often directed by humans who decide which pairs will mate to increase the chances of offspring having certain desirable traits. This is known as <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1501\"><strong>a<\/strong><strong>rtificial <\/strong><strong>s<\/strong><strong>election<\/strong><em>.<\/em><\/a><\/p>\n<p class=\"import-Normal\">Among humans, in addition to phenotypic traits, cultural traits such as religion and ethnicity may also influence assortative mating patterns.<\/p>\n<h3 class=\"import-Normal\"><strong>Defining a Species<\/strong><\/h3>\n<p class=\"import-Normal\"><em>Species<\/em> are organisms whose individuals are capable of breeding because they are biologically and behaviorally compatible to produce viable, fertile offspring. <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1502\"><strong>Viable offspring<\/strong><\/a> are those offspring that are healthy enough to survive to adulthood. <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1503\"><strong>Fertile offspring<\/strong><\/a> are able to reproduce successfully, resulting in offspring of their own. Both conditions must be met for individuals to be considered part of the same species. As you can imagine, these criteria complicate the identification of distinct species in fossilized remains of extinct populations. In those cases, we must examine how much phenotypic variation is typically found within a comparable modern-day species; we can then determine whether the fossilized remains fall within the expected range of variation for a single species.<\/p>\n<p class=\"import-Normal\">Some species have subpopulations that are regionally distinct. These are classified as separate <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1505\">subspecies<\/a><\/strong> because they have their own unique phenotypes and are geographically isolated from one another. However, if they do happen to encounter one another, they are still capable of successful interbreeding.<\/p>\n<p class=\"import-Normal\">There are many examples of sterile hybrids that are offspring of parents from two different species. For example, horses and donkeys can breed and have offspring together. Depending on which species is the mother and which is the father, the offspring are either called mules, or hennies. Mules and hennies can live full life spans but are not able to have offspring of their own. Likewise, tigers and lions have been known to mate and have viable offspring. Again, depending on which species is the mother and which is the father, these offspring are called either ligers or tigons. Like mules and hennies, ligers and tigons are unable to reproduce. In each of these cases, the mismatched set of chromosomes that the offspring inherit produce an adequate set of functioning genes for the hybrid offspring; however, once mixed and divided in meiosis, the gametes don\u2019t contain the full complement of genes needed for survival in the third generation.<\/p>\n<h3 class=\"import-Normal\"><strong>Micro- to Macroevolution<\/strong><\/h3>\n<p class=\"import-Normal\"><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1506\"><strong>Microevolution<\/strong><\/a> refers to changes in allele frequencies within breeding populations\u2014that is, within single species. <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1507\"><strong>Macroevolution<\/strong><\/a> describes how the similarities and differences between species, as well as the phylogenetic relationships with other taxa, lead to changes that result in the emergence of new species. Consider our example of the peppered moth that illustrated microevolution over time, via directional selection favoring the peppered allele when the trees were clean and the dark pigment allele when the trees were sooty. Imagine that environmental regulations had cleaned up the air pollution in one part of the nation, while the coal-fired factories continued to spew soot in another area. If this went on long enough, it\u2019s possible that two distinct moth populations would eventually emerge\u2014one containing only the peppered allele and the other only harboring the dark pigment allele.<\/p>\n<p class=\"import-Normal\">When a single population divides into two or more separate species, it is called <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1508\"><strong>speciation<\/strong><\/a>. The changes that prevent successful breeding between individuals who descended from the same ancestral population may involve chromosomal rearrangements, changes in the ability of the sperm from one species to permeate the egg membrane of the other species, or dramatic changes in hormonal schedules or mating behaviors that prevent members from the new species from being able to effectively pair up.<\/p>\n<p class=\"import-Normal\">There are two types of speciation: allopatric and sympatric. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1509\">Allopatric speciation<\/a><\/strong> is caused by long-term <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1510\"><strong>isolation<\/strong><\/a> (physical separation) of subgroups of the population (Figure 5.22). Something occurs in the environment\u2014perhaps a river changes its course and splits the group, preventing them from breeding with members on the opposite riverbank. Over many generations, new mutations and adaptations to the different environments on each side of the river may drive the two subpopulations to change so much that they can no longer produce fertile, viable offspring, even if the barrier is someday removed.<\/p>\n<figure style=\"width: 1000px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image23-2.png\" alt=\"Process of isolation leading to speciation, as described in the figure caption.\" width=\"1000\" height=\"379\" \/><figcaption class=\"wp-caption-text\">Figure 5.22: Isolation leading to speciation: a. original population before isolation; b. a barrier divides the population and prevents interbreeding between the two groups; c. time passes, and the populations become genetically distinct; d. after many generations, the two populations are no longer biologically or behaviorally compatible, thus can no longer interbreed, even if the barrier is removed. Credit: <a class=\"rId121\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Isolation Leading to Speciation (Figure 4.19)<\/a> original to <a class=\"rId122\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a class=\"rId123\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\"><strong>Sympatric speciation<\/strong> occurs when the population splits into two or more separate species while remaining located together <em>without<\/em> a physical barrier. This typically results from a new mutation that pops up among some members of the population that prevents them from successfully reproducing with anyone who does not carry the same mutation. This is seen particularly often in plants, as they have a higher frequency of chromosomal duplications.<\/p>\n<p class=\"import-Normal\">One of the quickest rates of speciation is observed in the case of adaptive radiation. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1512\">Adaptive radiation<\/a><\/strong> refers to the situation in which subgroups of a single species rapidly diversify and adapt to fill a variety of ecological niches. An <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1513\"><strong>e<\/strong><strong>cological niche<\/strong><\/a> is a set of constraints and resources that is available in an environmental setting. Evidence for adaptive radiations is often seen after population bottlenecks. A mass disaster kills off many species, and the survivors have access to a new set of territories and resources that were either unavailable or much coveted and fought over before the disaster. The offspring of the surviving population will often split into multiple species, each of which stems from members in that first group of survivors who happened to carry alleles that were advantageous for a particular niche.<\/p>\n<p class=\"import-Normal\">The classic example of adaptive radiation brings us back to Charles Darwin and his observations of the many species of finches on the Galapagos Islands. We are still not sure how the ancestral population of finches first arrived on that remote Pacific Island chain, but they found themselves in an environment filled with various insects, large and tiny seeds, fruit, and delicious varieties of cactus. Some members of that initial population carried alleles that gave them advantages for each of these dietary niches. In subsequent generations, others developed new mutations, some of which were beneficial. These traits were selected for, making the advantageous alleles more common among their offspring. As the finches spread from one island to the next, they would be far more likely to find mates among the birds on their new island. Birds feeding in the same area were then more likely to mate together than birds who have different diets, contributing to additional assortative mating. Together, these evolutionary mechanisms caused rapid speciation that allowed the new species to make the most of the various dietary niches (Figure 5.23).<\/p>\n<figure style=\"width: 619px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image24-1.png\" alt=\"A family tree of finches with different sized beaks.\" width=\"619\" height=\"325\" \/><figcaption class=\"wp-caption-text\">Figure 5.23: Darwin\u2019s finches demonstrating Adaptive Radiation. Credit: <a class=\"rId125\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-3\/\">Darwin\u2019s finches (Figure 4.20)<\/a> original to <a class=\"rId126\" href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson is under a <a class=\"rId127\" href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">In today\u2019s modern world, understanding these evolutionary processes is crucial for developing immunizations and antibiotics that can keep up with the rapid mutation rate of viruses and bacteria. This is also relevant to our food supply, which relies, in large part, on the development of herbicides and pesticides that keep up with the mutation rates of pests and weeds. Viruses, bacteria, agricultural pests, and weeds have all shown great flexibility in developing alleles that make them resistant to the latest medical treatment, pesticide, or herbicide. Billion-dollar industries have specialized in trying to keep our species one step ahead of the next mutation in the pests and infectious diseases that put our survival at risk.<\/p>\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Review Questions<strong><br \/>\n<\/strong><\/h2>\n<ul>\n<li>Summarize the Modern Synthesis and provide several examples of how it is relevant to questions and problems in our world today.<\/li>\n<li>You inherit a house from a long-lost relative that contains a fancy aquarium, filled with a variety of snails. The phenotypes include large snails and small snails; red, black, and yellow snails; and solid, striped, and spotted snails. Devise a series of experiments that would help you determine how many snail species are present in your aquarium.<\/li>\n<li>Match the correct force of evolution with the correct real-world example:<br \/>\na. Mutationi. 5-alpha reductase deficiency<br \/>\nb. Genetic Driftii. Peppered Moths<br \/>\nc. Gene Flowiii. Neurofibromatosis Type 1<br \/>\nd. Natural Selectioniv. Scutellata Honey Bees<\/li>\n<li>Imagine a population of common house mice (<em>Mus musculus<\/em>). Draw a comic strip illustrating how mutation, genetic drift, gene flow, and natural selection might transform this population over several (or more) generations.<\/li>\n<li>\n<p class=\"import-Normal\">The many breeds of the single species of domestic dog (<em>Canis<\/em> <em>familiaris<\/em>) provide an extreme example of microevolution. Discuss why this is the case. What future scenarios can you imagine that could potentially transform the domestic dog into an example of macroevolution?<\/p>\n<\/li>\n<li>\n<p class=\"import-Normal\">The ability to roll one\u2019s tongue (lift the outer edges of the tongue to touch each other, forming a tube) is a dominant trait. In a small town of 1,500 people, 500 can roll their tongues. Use the Hardy-Weinberg formula to determine how many individuals in the town are homozygous dominant, heterozygous, and homozygous recessive.<\/p>\n<\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\">Key Terms<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\"><strong>5-alpha reductase deficiency<\/strong>: An autosomal recessive syndrome that manifests when a child having both X and Y sex chromosomes inherits two nonfunctional (mutated) copies of the SRD5A2 gene, producing a deficiency in a hormone necessary for development in infancy of typical male genitalia. These children often appear at birth to have female genitalia, but they develop a penis and other sexual characteristics when other hormones kick in during puberty.<\/p>\n<p class=\"import-Normal\"><strong>Adaptive radiation<\/strong>: The situation in which subgroups of a single species rapidly diversify and adapt to fill a variety of ecological niches.<\/p>\n<p class=\"import-Normal\"><strong>Admixture<\/strong>: A term often used to describe gene flow between human populations. Sometimes also used to describe gene flow between nonhuman populations.<\/p>\n<p class=\"import-Normal\"><strong>Allele frequency<\/strong>: The ratio, or percentage, of one allele compared to the other alleles for that gene within the study population.<\/p>\n<p class=\"import-Normal\"><strong>Alleles<\/strong>: Variant forms of genes.<\/p>\n<p class=\"import-Normal\"><strong>Allopatric speciation<\/strong>: Speciation caused by long-term isolation (physical separation) of subgroups of the population.<\/p>\n<p class=\"import-Normal\"><strong>Antibiotics<\/strong>: Medicines prescribed to treat bacterial infections.<\/p>\n<p class=\"import-Normal\"><strong>Artificial selection<\/strong>: Human-directed assortative mating among domestic animals, such as pets and livestock, designed to increase the chances of offspring having certain desirable traits.<\/p>\n<p class=\"import-Normal\"><strong>Asexual reproduction<\/strong>: Reproduction via mitosis, whereby offspring are clones of the parents.<\/p>\n<p class=\"import-Normal\"><strong>Autosomal dominant<\/strong>: A phenotype produced by a gene on an autosomal chromosome that is expressed, to the exclusion of the recessive phenotype, in heterozygotes.<\/p>\n<p class=\"import-Normal\"><strong>Autosomal recessive<\/strong>: A phenotype produced by a gene on an autosomal chromosome that is expressed only in individuals homozygous for the recessive allele.<\/p>\n<p class=\"import-Normal\"><strong>Balanced translocations<\/strong>: Chromosomal translocations in which the genes are swapped but no genetic information is lost.<\/p>\n<p class=\"import-Normal\"><strong>Balancing selection<\/strong>: A pattern of natural selection that occurs when the extremes of a trait are selected against, favoring the intermediate phenotype (a.k.a. stabilizing selection).<\/p>\n<p class=\"import-Normal\"><strong>Beneficial mutations<\/strong>: Mutations that produce some sort of an advantage to the individual.<\/p>\n<p class=\"import-Normal\"><strong>Benign<\/strong>: Noncancerous. Benign tumors may cause problems due to the area in which they are located (e.g., they might put pressure on a nerve or brain area), but they will not release cells that aggressively spread to other areas of the body.<\/p>\n<p class=\"import-Normal\"><strong>Caf\u00e9-au-lait spots (CALS)<\/strong>: Flat, brown birthmark-like spots on the skin, commonly associated with Neurofibromatosis Type 1.<\/p>\n<p class=\"import-Normal\"><strong>Chromosomal translocations<\/strong>: The transfer of DNA between nonhomologous chromosomes.<\/p>\n<p class=\"import-Normal\"><strong>Chromosomes<\/strong>: Molecules that carry collections of genes.<\/p>\n<p class=\"import-Normal\"><strong>Codons<\/strong>: Three-nucleotide units of DNA that function as three-letter \u201cwords,\u201d encoding instructions for the addition of one amino acid to a protein or indicating that the protein is complete.<\/p>\n<p class=\"import-Normal\"><strong>Cretaceous\u2013Paleogene extinction<\/strong>: A mass disaster caused by an asteroid that struck the earth approximately 66 million years ago and killed 75% of life on Earth, including all terrestrial dinosaurs. (a.k.a. K-Pg Extinction, Cretatious-Tertiary Extinction, and K-T Extinction).<\/p>\n<p class=\"import-Normal\"><strong>Crossover events<\/strong>: Chromosomal alterations that occur when DNA is swapped between homologous chromosomes while they are paired up during meiosis I.<\/p>\n<p class=\"import-Normal\"><strong>Cutaneous neurofibromas<\/strong>: Neurofibromas that manifest as spherical bumps on or just under the surface of the skin.<\/p>\n<p class=\"import-Normal\"><strong>Deleterious mutation<\/strong>: A mutation producing negative effects to the individual such as the beginnings of cancers or heritable disorders.<\/p>\n<p class=\"import-Normal\"><strong>Deletions<\/strong>: Mutations that involve the removal of one or more nucleotides from a DNA sequence.<\/p>\n<p class=\"import-Normal\"><strong>Derivative chromosomes<\/strong>: New chromosomal structures resulting from translocations.<\/p>\n<p class=\"import-Normal\"><strong><em>Dictyostelium discoideum<\/em><\/strong>: A species of social amoebae that has been widely used for laboratory research. Laboratory strains of <em>Dictyostelium discoideum <\/em>all carry mutations in the <em>NF1<\/em> gene, which is what allows them to survive on liquid media (agar) in Petri dishes.<\/p>\n<p class=\"import-Normal\"><strong>Directional selection<\/strong>: A pattern of natural selection in which one phenotype is favored over the other, causing the frequencies of the associated advantageous alleles to gradually increase.<\/p>\n<p class=\"import-Normal\"><strong>Disruptive selection<\/strong>: A pattern of natural selection that occurs when both extremes of a trait are advantageous and intermediate phenotypes are selected against (a.k.a. diversifying selection).<\/p>\n<p class=\"import-Normal\"><strong>DNA repair mechanisms<\/strong>: Enzymes that patrol and repair DNA in living cells.<\/p>\n<p class=\"import-Normal\"><strong>DNA transposons<\/strong>: Transposons that are clipped out of the DNA sequence itself and inserted elsewhere in the genome.<\/p>\n<p class=\"import-Normal\"><strong>Ecological niche<\/strong>: A set of constraints and resources that are available in an environmental setting.<\/p>\n<p class=\"import-Normal\"><strong>Ellis-van Creveld syndrome<\/strong>: An autosomal recessive disorder characterized by short stature (dwarfism), polydactyly (the development of more than five digits [fingers or toes] on the hands or feet), abnormal tooth development, and heart defects. Estimated to affect approximately one in 60,000 individuals worldwide, among the Old Order Amish of Lancaster County, the rate is estimated to be as high as one in every 200 births.<\/p>\n<p class=\"import-Normal\"><strong>Evolution<\/strong>: A change in the allele frequencies in a population over time.<\/p>\n<p class=\"import-Normal\"><strong>Exons<\/strong>: The DNA sequences within a gene that directly encode protein sequences. After being transcribed into messenger RNA, the introns (DNA sequences within a gene that do not directly encode protein sequences) are clipped out, and the exons are pasted together prior to translation.<\/p>\n<p class=\"import-Normal\"><strong>Fertile offspring<\/strong>: Offspring that can successfully reproduce, resulting in offspring of their own.<\/p>\n<p class=\"import-Normal\"><strong>Founder effect<\/strong>: A type of genetic drift that occurs when members of a population leave the main or \u201cparent\u201d group and form a new population that no longer interbreeds with the other members of the original group.<\/p>\n<p class=\"import-Normal\"><strong>Frameshift mutations<\/strong>: Types of indels that involve the insertion or deletion of any number of nucleotides that is not a multiple of three. These \u201cshift the reading frame\u201d and cause all codons beyond the mutation to be misread.<\/p>\n<p class=\"import-Normal\"><strong>Gametes<\/strong>: The reproductive cells, produced through meiosis (a.k.a. germ cells or sperm or egg cells).<\/p>\n<p class=\"import-Normal\"><strong>Gene<\/strong>: A sequence of DNA that provides coding information for the construction of proteins.<\/p>\n<p class=\"import-Normal\"><strong>Gene flow<\/strong>: The movement of alleles from one population to another. This is one of the forces of evolution.<\/p>\n<p class=\"import-Normal\"><strong>Gene pool<\/strong>: The entire collection of genetic material in a breeding community that can be passed on from one generation to the next.<\/p>\n<p class=\"import-Normal\"><strong>Genetic drift<\/strong>: Random changes in allele frequencies within a population from one generation to the next. This is one of the forces of evolution.<\/p>\n<p class=\"import-Normal\"><strong>Genotype<\/strong>: The set of alleles that an individual has for a given gene.<\/p>\n<p class=\"import-Normal\"><strong>Genotype frequencies<\/strong>: The ratios or percentages of the different homozygous and heterozygous genotypes in the population.<\/p>\n<p class=\"import-Normal\"><strong><em>Guevedoces<\/em><\/strong>: The term coined locally in the Dominican Republic for the condition scientifically known as 5-alpha reductase deficiency. The literal translation is \u201cpenis at twelve.\u201d<\/p>\n<p class=\"import-Normal\"><strong>Hardy-Weinberg Equilibrium<\/strong>: A mathematical formula (<em>1=p<\/em><sup><em>2<\/em><\/sup><em> + 2pq + q<\/em><sup><em>2<\/em><\/sup> ) that allows estimation of the number and distribution of dominant and recessive alleles in a population.<\/p>\n<p class=\"import-Normal\"><strong>Harlequin ladybeetle<\/strong>: A species of ladybeetle, native to East Asia, that was introduced to Europe and the Americas as a form of pest control. After many decades of use, one of the North American strains developed the ability to reproduce in diverse environments, causing it to spread rapidly throughout the Americas, Europe, and Africa. It has hybridized with European strains and is now a major pest in its own right.<\/p>\n<p class=\"import-Normal\"><strong>Heterozygous genotype<\/strong>: A genotype comprising two different alleles.<\/p>\n<p class=\"import-Normal\"><strong>Homozygous genotype<\/strong>: A genotype comprising an identical set of alleles.<\/p>\n<p class=\"import-Normal\"><strong>Hybridization<\/strong>: A term often used to describe gene flow between nonhuman populations.<\/p>\n<p class=\"import-Normal\"><strong>Inbreeding<\/strong>: The selection of mates exclusively from within a small, closed population.<\/p>\n<p class=\"import-Normal\"><strong>Indels<\/strong>: A class of mutations that includes both insertions and deletions.<\/p>\n<p class=\"import-Normal\"><strong>Inherited mutation<\/strong>: A mutation that has been passed from parent to offspring.<\/p>\n<p class=\"import-Normal\"><strong>Insertions<\/strong>: Mutations that involve the addition of one or more nucleotides into a DNA sequence.<\/p>\n<p class=\"import-Normal\"><strong>Isolation<\/strong>: Prevention of a population subgroup from breeding with other members of the same species due to a physical barrier or, in humans, a cultural rule.<\/p>\n<p class=\"import-Normal\"><strong>Last Universal Common Ancestor (LUCA)<\/strong>: The ancient organism from which all living things on Earth are descended.<\/p>\n<p class=\"import-Normal\"><strong>Macroevolution<\/strong>: Changes that result in the emergence of new species, how the similarities and differences between species, as well as the phylogenetic relationships with other taxa, lead to changes that result in the emergence of new species.<\/p>\n<p class=\"import-Normal\"><strong>Malaria<\/strong>: A frequently deadly mosquito-borne disease caused by infection of the blood by a <em>Plasmodium<\/em> parasite.<\/p>\n<p class=\"import-Normal\"><strong>Malignant<\/strong>: Cancerous. Malignant tumors grow aggressively and their cells may metastasize (travel through the blood or lymph systems) to form new, aggressive tumors in other areas of the body.<\/p>\n<p class=\"import-Normal\"><strong>Microevolution<\/strong>: Changes in allele frequencies within breeding populations\u2014that is, within a single species.<\/p>\n<p class=\"import-Normal\"><strong>Modern Synthesis<\/strong>: The integration of Darwin\u2019s, Mendel\u2019s, and subsequent research into a unified theory of evolution.<\/p>\n<p class=\"import-Normal\"><strong>Monosomies<\/strong>: Conditions resulting from a nondisjunction event, in which a cell ends up with only one copy of a chromosome. In humans, a single X chromosome is the only survivable monosomy.<\/p>\n<p class=\"import-Normal\"><strong>Mutation<\/strong>: A change in the nucleotide sequence of the genetic code. This is one of the forces of evolution.<\/p>\n<p class=\"import-Normal\"><strong>Natural selection<\/strong>: An evolutionary process that occurs when certain phenotypes confer an advantage or disadvantage in survival and\/or reproductive success. This is one of the forces of evolution, and it was first identified by Charles Darwin.<\/p>\n<p class=\"import-Normal\"><strong>Negative assortative mating<\/strong>: A pattern that occurs when individuals tend to select mates with qualities different from their own.<\/p>\n<p class=\"import-Normal\"><strong>Neurofibromas<\/strong>: Nerve sheath tumors that are common symptoms of Neurofibromatosis Type 1.<\/p>\n<p class=\"import-Normal\"><strong>Neurofibromatosis Type 1<\/strong>: An autosomal dominant genetic disorder affecting one in every 3,000 people. It is caused by mutation of the <em>NF1<\/em> gene on Chromosome 17, resulting in a defective neurofibromin protein. The disorder is characterized by neurofibromas, caf\u00e9-au-lait spots, and a host of other potential symptoms.<\/p>\n<p class=\"import-Normal\"><strong>NF1<\/strong>: An abbreviation for Neurofibromatosis Type 1. When italicized, <em>NF1 <\/em>refers to the gene on Chromosome 17 that encodes the neurofibromin protein.<\/p>\n<p class=\"import-Normal\"><strong>Nondisjunction events<\/strong>: Chromosomal abnormalities that occur when the homologous chromosomes (in meiosis I) or sister chromatids (in meiosis II and mitosis) fail to separate after pairing. The result is that both chromosomes or chromatids end up in the same daughter cell, leaving the other daughter cell without any copy of that chromosome.<\/p>\n<p class=\"import-Normal\"><strong>Nonrandom mating<\/strong>: A scenario in which mate choice within a population follows a nonrandom pattern (a.k.a. assortative mating).<\/p>\n<p class=\"import-Normal\"><strong>Nonsynonymous mutation<\/strong>: A point mutation that causes a change in the resulting protein.<\/p>\n<p class=\"import-Normal\"><strong>Old Order Amish<\/strong>: A culturally isolated population in Lancaster County, Pennsylvania, that has approximately 50,000 current members, all of whom can trace their ancestry back to a group of approximately eighty individuals. This group has high rates of certain genetics disorders, including Ellis-van Creveld syndrome.<\/p>\n<p class=\"import-Normal\"><strong>Origins of life<\/strong>: How the first living organism came into being.<\/p>\n<p class=\"import-Normal\"><strong>Peacock<\/strong>: The male sex of the peafowl, famous for its large, colorful tail, which it dramatically displays to attract mates. (The female of the species is known as a peahen.)<\/p>\n<p class=\"import-Normal\"><strong>Peppered moth<\/strong>: A species of moth (<em>Biston betularia<\/em>) found in England that has light and dark phenotypes. During the Industrial Revolution, when soot blackened the trees, the frequency of the previously rare dark phenotype dramatically increased, as lighter-colored moths were easier for birds to spot against the sooty trees. After environmental regulations eliminated the soot, the lighter-colored phenotype gradually became most common again.<\/p>\n<p class=\"import-Normal\"><strong>Phenotype<\/strong>: The observable traits that are produced by a genotype.<\/p>\n<p class=\"import-Normal\"><strong>Phylogenetic tree of life<\/strong>: A family tree of all living organisms, based on genetic relationships.<\/p>\n<p class=\"import-Normal\"><strong>Phylogenies<\/strong>: Genetically determined family lineages.<\/p>\n<p class=\"import-Normal\"><strong><em>Plasmodium<\/em><\/strong>: A genus of mosquito-borne parasite. Several <em>Plasmodium<\/em> species cause malaria when introduced to the human bloodstream via a mosquito bite.<\/p>\n<p class=\"import-Normal\"><strong>Plexiform neurofibromas<\/strong>: Neurofibromas that involve whole branches of nerves, often giving the appearance that the surface of the skin is \u201cmelting.\u201d<\/p>\n<p class=\"import-Normal\"><strong>Point mutation<\/strong>: A single-letter (single-nucleotide) change in the genetic code, resulting in the substitution of one nucleic acid base for a different one.<\/p>\n<p class=\"import-Normal\"><strong>Polymorphisms<\/strong>: Multiple forms of a trait; alternative phenotypes within a given species.<\/p>\n<p class=\"import-Normal\"><strong>Population<\/strong>: A group of individuals who are genetically similar enough and geographically near enough to one another that they can breed and produce new generations of individuals.<\/p>\n<p class=\"import-Normal\"><strong>Population bottleneck<\/strong>: A type of genetic drift that occurs when the number of individuals in a population drops dramatically due to some random event.<\/p>\n<p class=\"import-Normal\"><strong>Positive assortative mating<\/strong>: A pattern that results from a tendency for individuals to mate with others who share similar phenotypes.<\/p>\n<p class=\"import-Normal\"><strong>Retrotransposons<\/strong>: Transposons that are transcribed from DNA into RNA, and then are \u201creverse transcribed,\u201d to insert the copied sequence into a new location in the DNA.<\/p>\n<p class=\"import-Normal\"><strong>Scutellata honey bees<\/strong>: A strain of honey bees that resulted from the hybridization of African and European honey bee subspecies. These bees were accidentally released into the wild in 1957 in Brazil and have since spread throughout South and Central America and into the United States. Also known as \u201ckiller bees,\u201d they tend to be very aggressive in defense of their hives and have caused many fatal injuries to humans and livestock.<\/p>\n<p class=\"import-Normal\"><strong>Sexual reproduction<\/strong>: Reproduction via meiosis and combination of gametes. Offspring inherit genetic material from both parents.<\/p>\n<p class=\"import-Normal\"><strong>Sexual selection<\/strong>: An aspect of natural selection in which the selective pressure specifically affects reproductive success (the ability to successfully breed and raise offspring).<\/p>\n<p class=\"import-Normal\"><strong>Sickle cell anemia<\/strong>: An autosomal recessive genetic disorder that affects millions of people worldwide. It is most common in Africa, countries around the Mediterranean Sea, and eastward as far as India. Homozygotes for the recessive allele develop the disorder, which produce misshapen red blood cells that cause iron deficiency, painful episodes of oxygen-deprivation in localized tissues, and a host of other symptoms. In heterozygotes, though, the sickle cell allele confers a greater resistance to malaria.<\/p>\n<p class=\"import-Normal\"><strong>Somatic cells<\/strong>: The cells of our organs and other body tissues (all cells except gametes) that replicate by mitosis.<\/p>\n<p class=\"import-Normal\"><strong>Speciation<\/strong>: The process by which a single population divides into two or more separate species.<\/p>\n<p class=\"import-Normal\"><strong>Species<\/strong>: Organisms whose individuals are capable of breeding because they are biologically and behaviorally compatible to produce viable, fertile offspring.<\/p>\n<p class=\"import-Normal\"><strong>Spontaneous mutation<\/strong>: A mutation that occurs due to random chance or unintentional exposure to mutagens. In families, a spontaneous mutation is the first case, as opposed to mutations that are inherited from parents.<\/p>\n<p class=\"import-Normal\"><strong>Subspecies<\/strong>: A distinct subtype of a species. Most often, this is a geographically isolated population with unique phenotypes; however, it remains biologically and behaviorally capable of interbreeding with other populations of the same species.<\/p>\n<p class=\"import-Normal\"><strong>Sympatric speciation<\/strong>: When a population splits into two or more separate species while remaining located together without a physical (or cultural) barrier.<\/p>\n<p class=\"import-Normal\"><strong>Synonymous mutation<\/strong>: A point mutation that does not change the resulting protein.<\/p>\n<p class=\"import-Normal\"><strong>Transposable elements<\/strong>: Fragments of DNA that can \u201cjump\u201d around in the genome.<\/p>\n<p class=\"import-Normal\"><strong>Transposon<\/strong>: Another term for \u201ctransposable element.\u201d<\/p>\n<p class=\"import-Normal\"><strong>Trisomies<\/strong>: Conditions in which three copies of the same chromosome end up in a cell, resulting from a nondisjunction event. Down syndrome, Edwards syndrome, and Patau syndrome are trisomies.<\/p>\n<p class=\"import-Normal\"><strong>Unbalanced translocations<\/strong>: Chromosomal translocations in which there is an unequal exchange of genetic material, resulting in duplication or loss of genes.<\/p>\n<p class=\"import-Normal\"><strong>UV crosslinking<\/strong>: A type of mutation in which adjacent thymine bases bind to one another in the presence of UV light.<\/p>\n<p class=\"import-Normal\"><strong>Viable offspring<\/strong>: Offspring that are healthy enough to survive to adulthood.<\/p>\n<p class=\"import-Normal\"><strong>Xeroderma pigmentosum<\/strong>: An autosomal recessive disease in which DNA repair mechanisms do not function correctly, resulting in a host of problems especially related to sun exposure, including severe sunburns, dry skin, heavy freckling, and other pigment changes.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration<\/h2>\n<p>Explore Evolution on <a href=\"https:\/\/www.hhmi.org\/biointeractive\/evolution-collection\">HHMI\u2019s Biointeractive website<\/a>.<\/p>\n<p>Teaching Evolution through <a href=\"https:\/\/humanorigins.si.edu\/education\/teaching-evolution-through-human-examples\">Human Examples, Smithsonian Museum of Natural History websites<\/a>.<\/p>\n<h2 class=\"import-Normal\">References<\/h2>\n<p class=\"import-Normal\">Bloomfield, Gareth, David Traynor, Sophia P. Sander, Douwe M. Veltman, Justin A. Pachebat, and Robert R. Kay. 2015. \u201cNeurofibromin Controls Macropinocytosis and Phagocytosis in <em>Dictyostelium<\/em>.\u201d <em>eLife<\/em> 4:e04940.<\/p>\n<p class=\"import-Normal\">Chaix, Rapha\u00eblle, Chen Cao, and Peter Donnelly. 2008. \u201cIs Mate Choice in Humans MHC-Dependent?\u201d\u00a0<em>PLoS Genetics<\/em>\u00a04 (9): e1000184.<\/p>\n<p class=\"import-Normal\">Cook, Laurence\u00a0M. 2003. \"The Rise and Fall of the\u00a0<em>Carbonaria<\/em>\u00a0Form of the Peppered Moth.\" <em>The Quarterly Review of Biology<\/em> 78 (4): 399\u2013417.<\/p>\n<p class=\"import-Normal\">Cota, Bruno C\u00e9zar Lage, Jo\u00e3o Gabriel Marques Fonseca, Luiz Oswaldo Carneiro Rodrigues, Nilton Alves de Rezende, Pollyanna Barros Batista, Vincent Michael Riccardi, and Luciana Macedo de Resende. 2018. \u201cAmusia and Its Electrophysiological Correlates in Neurofibromatosis Type 1.\u201d <em>Arquivos de Neuro-Psiquiatria<\/em> 76 (5): 287\u2013295.<\/p>\n<p class=\"import-Normal\">D\u2019Asdia, Maria Cecilia, Isabella Torrente, Federica Consoli, Rosangela Ferese, Monia Magliozzi, Laura Bernardini, Valentina Guida, et al. 2013. \u201cNovel and Recurrent EVC and EVC2 Mutations in Ellis-van Creveld Syndrome and Weyers Acrofacial Dyostosis.\u201d <em>European Journal of Medical Genetics<\/em> 56 (2): 80\u201387.<\/p>\n<p class=\"import-Normal\">Dobzhansky, Theodosius. 1937. <em>Genetics and the Origin of Species. <\/em>Columbia University Biological Series. New York: Columbia University Press.<\/p>\n<p class=\"import-Normal\">Facon, Beno\u00eet, Laurent Crespin, Anne Loiseau, Eric Lombaert, Alexandra Magro, and Arnaud Estoup. 2011. \u201cCan Things Get Worse When an Invasive Species Hybridizes? The Harlequin Ladybird\u00a0<em>Harmonia axyridis<\/em>\u00a0in France as a Case Study.\u201d\u00a0<em>Evolutionary Applications<\/em> 4 (1): 71\u201388.<\/p>\n<p class=\"import-Normal\">Fisher, Ronald A. 1919. \"The Correlation between Relatives on the Supposition of Mendelian Inheritance.\" <em>Transactions of the Royal Society of Edinburgh<\/em> 52 (2): 399\u2013433.<\/p>\n<p class=\"import-Normal\">Ford, E. B. 1942.\u00a0<em>Genetics for Medical Students<\/em>. London: Methuen.<\/p>\n<p class=\"import-Normal\" style=\"background-color: #ffffff\">Ford, E. B.\u00a01949.\u00a0<em>Mendelism and Evolution<\/em>. London: Methuen.<\/p>\n<p class=\"import-Normal\">Grant, Bruce S. 1999. \u201cFine-tuning the Peppered Moth Paradigm.\u201d <em>Evolution<\/em> 53 (3): 980\u2013984.<\/p>\n<p class=\"import-Normal\">Haldane, J. B. S.\u00a01924.\u00a0\u201cA Mathematical Theory of Natural and Artificial Selection (Part 1).\u201d <em>Transactions of the Cambridge Philosophical Society<\/em>\u00a023 (2):19\u201341.<\/p>\n<p>Hoelzel, A. R., Gkafas, G. A., Kang, H., Sarigol, F., Le Boeuf, B., Costa, D. P., Beltran, R. S., Reiter, J., Robinson, P. W., McInerney, N., Seim, I., Sun, S., Fan, G., &amp; Li, S. (2024). Genomics of post-bottleneck recovery in the northern elephant seal. Nature Ecology &amp; Evolution, 8, 686\u2013694. https:\/\/doi.org\/10.1038\/s41559-024-02337-4<\/p>\n<p class=\"import-Normal\">Imperato-McGinley, J., and Y.-S. Zhu. 2002. \u201cAndrogens and Male Physiology: The Syndrome of 5 Alpha-Reductase-2 Deficiency.\u201d\u00a0<em>Molecular and Cellular Endocrinology <\/em>198 (1-2): 51\u201359.<\/p>\n<p class=\"import-Normal\">Jablonski, David, and W. G. Chaloner. 1994. \"Extinctions in the Fossil Record.\u201d\u00a0<em>Philosophical Transactions of the Royal Society of London\u00a0B: Biological Sciences<\/em>\u00a0344 (1307): 11\u201317.<\/p>\n<p class=\"import-Normal\">Livi-Bacci, Massimo. 2006. \u201cThe Depopulation of Hispanic America after the Conquest.\u201d <em>Population Development and Review<\/em> 32 (2): 199\u2013232.<\/p>\n<p class=\"import-Normal\">Lombaert, Eric, Thomas Guillemaud, Jean-Marie Cornuet, Thibaut Malausa, Beno\u00eet Facon, and Arnaud Estoup. 2010. \"Bridgehead Effect in the Worldwide Invasion of the Biocontrol Harlequin Ladybird.\u201d <em>PLoS ONE<\/em> 5 (3): e9743.<\/p>\n<p class=\"import-Normal\">Martins, Aline Stangherlin, Ann Kristine Jansen, Luiz Oswaldo Carneiro Rodrigues, Camila Maria Matos, Marcio Leandro Ribeiro Souza, Juliana Ferreira de Souza, Maria de F\u00e1tima Haueisen Sander Diniz, et al. 2016. \u201cLower Fasting Blood Glucose in Neurofibromatosis Type 1.\u201d <em>Endocrine Connections<\/em> 5 (1): 28\u201333.<\/p>\n<p class=\"import-Normal\">Pickering, Gary, James Lin, Roland Riesen, Andrew Reynolds, Ian Brindle, and George Soleas. 2004.\u00a0\"Influence of\u00a0<em>Harmonia axyridis<\/em>\u00a0on the Sensory Properties of White and Red Wine.\"\u00a0<em>American Journal of Enology and Viticulture<\/em>\u00a055 (2): 153\u2013159.<\/p>\n<p class=\"import-Normal\">Repunte-Canonigo Vez, Melissa A. Herman, Tomoya Kawamura, Henry R. Kranzler, Richard Sherva, Joel Gelernter, Lindsay A. Farrer, Marisa Roberto, and Pietro Paolo Sanna. 2015. \u201cNF1 Regulates Alcohol Dependence-Associated Excessive Drinking and Gamma-Aminobutyric Acid Release in the Central Amygdala in Mice and Is Associated with Alcohol Dependence in Humans.\u201d <em>Biological Psychiatry<\/em> 77 (10): 870\u2013879.<\/p>\n<p class=\"import-Normal\">Riccardi, Vincent M. 1992. <em>Neurofibromatosis: Phenotype, Natural History, and Pathogenesis.<\/em> Baltimore: Johns Hopkins University Press.<\/p>\n<p class=\"import-Normal\">Sanford, Malcolm T. 2006.\u00a0\"The Africanized Honey Bee in the Americas: A Biological Revolution with Human Cultural Implications, Part V\u2014Conclusion.\"\u00a0<em>American Bee Journal <\/em>146 (7): 597\u2013599.<\/p>\n<p class=\"import-Normal\">Sanna, Pietro Paolo, Cindy Simpson, Robert Lutjens, and George Koob. 2002. \u201cERK Regulation in Chronic Ethanol Exposure and Withdrawal.\u201d <em>Brain Research<\/em> 948 (1\u20132): 186\u2013191.<\/p>\n<p>Weber, DianaS., Stewart, B. S., Garza, J. Carlos., &amp; Lehman, N. (2000). An empirical genetic assessment of the severity of the northern elephant seal population bottleneck. Current Biology, 10(20), 1287\u20131290. https:\/\/doi.org\/10.1016\/s0960-9822(00)00759-4<\/p>\n<p class=\"import-Normal\">World Health Organization. 1996. \u201cControl of Hereditary Disorders: Report of WHO Scientific meeting (1996).\u201d WHO Technical Reports 865. Geneva: World Health Organization.<\/p>\n<p class=\"import-Normal\">World Health Organization. 2017. \u201cGlobal Priority List of Antibiotic-Resistant Bacteria to Guide Research, Discovery, and Development of New Antibiotics.\u201d Global Priority Pathogens List, February 27. Geneva: World Health Organization. https:\/\/www.who.int\/medicines\/publications\/WHO-PPL-Short_Summary_25Feb-ET_NM_WHO.pdf.<\/p>\n<p class=\"import-Normal\">Wright, Sewall. 1932. \"The Roles of Mutation, Inbreeding, Crossbreeding, and Selection in Evolution.\" <em>Proceedings of the Sixth International Congress on Genetics<\/em> 1 (6): 356\u2013366.<\/p>\n<h2 class=\"import-Normal\">Acknowledgment<strong><br \/>\n<\/strong><\/h2>\n<p class=\"import-Normal\">Many thanks to Dr. Vincent M. Riccardi for sharing his vast knowledge of neurofibromatosis and for encouraging me to explore it from an anthropological perspective.<\/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_1683_1708\"><div class=\"glossary__definition\" role=\"dialog\" data-id=\"term_1683_1708\"><div tabindex=\"-1\"><div class=\"__UNKNOWN__\">\n<div class=\"textbox\">\n<h6>Student contributors for this chapter from Salinda Hess' ANTH 203: <em>Emma Costa, Shima Gahima, Will Lefebvre, Audrey Ch\u00e9kina\u00ebl. <\/em><\/h6>\n<p>Learn more about <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/about-the-authors\/\">authors and editors<\/a>,<a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/student-contributors\/\"> Hess' student contributions<\/a>, and <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/versioning-history\/\">versioning history<\/a>.<\/p>\n<div class=\"textbox textbox--learning-objectives\">\n<header class=\"textbox__header\">\n<h2 class=\"textbox__title\">Learning Objectives<\/h2>\n<\/header>\n<div class=\"textbox__content\">\n<ul>\n<li class=\"import-Normal\">Describe how studying nonhuman primates is important in anthropology.<\/li>\n<li class=\"import-Normal\">Compare two ways of categorizing taxa: grades and clades.<\/li>\n<li class=\"import-Normal\">Define different types of traits used to evaluate primate taxa.<\/li>\n<li class=\"import-Normal\">Identify key ways that primates differ from other mammals.<\/li>\n<li class=\"import-Normal\">Distinguish between the major primate taxa using their key characteristics.<\/li>\n<li class=\"import-Normal\">Describe your place in nature by learning your taxonomic classification.<\/li>\n<\/ul>\n<\/div>\n<\/div>\n<p class=\"import-Normal\">You may be wondering why a field dedicated to the study of humans includes discussions of nonhuman animals. Our primary goal in biological anthropology is to understand how humans are similar to and different from the rest of the natural world, why we have the traits we have, and how we got to be the way we are. But to fully grasp our place in nature, we must look to our closest living relatives, the nonhuman primates. In this chapter, we focus on the organization and diversity within the Order Primates.<\/p>\n<h2 class=\"import-Normal\">Studying Primates in Biological Anthropology<\/h2>\n<p class=\"import-Normal\">Primates are one of at least twenty Orders belonging to the Class Mammalia, and probably one of the oldest. One genetic estimate puts the origin of primates at approximately 91 million years ago (mya), predating the extinction of the dinosaurs (Bininda-Emonds Et al. 2007). Today, the Order Primates is a diverse group of animals that includes lemurs and lorises, tarsiers, monkeys, apes, and humans, all of which are united in sharing a suite of anatomical, behavioural, and life history characteristics. While nonhuman primates are fascinating animals in their own right, their close relationship to humans makes them ideal for studying humans via <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_944\">homology<\/a>, <\/strong>looking at traits that are shared between taxa because they inherited the trait from a common ancestor. For example, humans (genus <em>Homo<\/em>) and chimpanzees (genus <em>Pan<\/em>) both share the trait of male cooperation in hunting. This trait\u2014along with many others that chimpanzees and humans share\u2014is likely homologous<em>, <\/em>meaning it was probably passed down from the last common ancestor of <em>Homo<\/em> and <em>Pan, <\/em>which lived about 6\u20138 million years ago.<\/p>\n<p class=\"import-Normal\">Nonhuman primates also make excellent comparators for learning about humans via <strong>analogy<\/strong>. Many nonhuman primates live in environments similar to those in which our ancestors lived and therefore exhibit traits similar to what we see in humans. For example, baboons and humans both have long legs. In humans, this is because about 1.7 million years ago, our ancestors moved into savanna habitats where longer legs helped them move more efficiently over long distances. Baboons, who also live in savanna habitats, independently evolved longer arms and legs for the same reason\u2014to be able to cover more ground, more efficiently. This means that having long legs is an analogous trait in baboons and humans: \u2014that is, this adaptation evolved independently in the two species but for the same purpose. Using homology and analogy, our closest living relatives provide the critical context in which to understand human biology, morphology, and behaviour. It is only by studying how humans compare with our primate relatives that we can fully comprehend our place in nature.<\/p>\n<h3 class=\"import-Normal\"><strong>Ways of Organizing Taxa<\/strong><\/h3>\n<p class=\"import-Normal\">You learned in Chapter 2 about Linnaeus and the hierarchical nature of taxonomic classification. Our goal in classifying taxa is to create categories that reflect clade relationships. A <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1116\">clade<\/a> <\/strong>is a grouping of organisms based on relatedness that reflects a branch of the evolutionary tree. Clade relationships are determined using traits shared by groups of taxa as well as genetic similarities. An example of a clade would be a grouping that includes humans, chimpanzees, bonobos, and gorillas (Figure 6.1). These taxa are in what is referred to as the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1118\">African clade<\/a><\/strong> of hominoids (a taxonomic group you will learn about later in this chapter). The African clade grouping reflects how humans, chimpanzees, bonobos, and gorillas all share a more recent ancestor with each other than any of them do with other species\u2014that is, we are on the same branch of the evolutionary tree. We know members of the African clade are most closely related based on shared morphological traits as well as genetic similarities. Excluded from this grouping is the orangutan, which is considered a member of the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1120\">Asian clade<\/a><\/strong> of hominoids.<\/p>\n<figure style=\"width: 800px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2023\/06\/image1-3.jpg\" alt=\"Diagram shows large-bodied hominoids grouped by grade or clade.\" width=\"800\" height=\"358\" \/><figcaption class=\"wp-caption-text\">Figure 6.1: Grades vs. Clades. A grade grouping of apes places orangutans, gorillas, chimpanzees, and bonobos together based on their similar appearance and lifestyle, but excludes humans. Clade classification is based on shared derived traits and genetic evidence (both reflecting close evolutionary relationships). A clade grouping of apes places humans with gorillas, chimpanzees, and bonobos., whereas orangutans are separated. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-4\/\">Grades vs. clades comparison (Figure 5.12)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Stephanie Etting 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:Orangutan_on_a_tree_(Unsplash).jpg\">Orangutan on a tree (Unsplash)<\/a> by Dawn Armfield, <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/legalcode\">public domain (CC0 1.0)<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Gorilla_Profile_(17997840570).jpg\">Gorilla Profile (17997840570)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/100915417@N07\">Charlie Marshall<\/a> from Bristol UK, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\">CC BY 2.0 License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Chimpanzee_(14679767561).jpg\">Chimpanzee (14679767561)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/120374925@N06\">Magnus Johansson<\/a>, modified (cropped), <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/legalcode\">CC BY-SA 2.0<\/a>; <a href=\"https:\/\/pixabay.com\/photos\/pointing-finger-hand-pointing-1922074\/\">Pointing finger (1922074)<\/a> by <a href=\"https:\/\/pixabay.com\/users\/truthseeker08-2411480\/\">truthseeker08<\/a>, <a href=\"https:\/\/pixabay.com\/service\/terms\/#license\">Pixabay License<\/a>.]<\/figcaption><\/figure>\n<p class=\"import-Normal\">In contrast, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1122\">grades<\/a> <\/strong>are groupings that reflect levels of adaptation or overall similarity and not necessarily evolutionary relationships. An example of a grade would be placing orangutans, gorillas, bonobos, and chimpanzees into a group, and excluding humans. Grouping in this way is based on the superficial similarities of the apes in being large-bodied, having lots of body hair, living in tropical forests, climbing and sleeping in trees, and so on. According to these criteria, humans seem to be unusual in that we differ in our morphology, behaviour, and ecology. Separating humans from the large-bodied apes is the system that was used historically. We now know that grouping orangutans, gorillas, bonobos, and chimpanzees and excluding humans does not accurately reflect our true evolutionary relationships. Since our goal in taxonomic classification is to organize animals to reflect their evolutionary relationships, we prefer to use clade classifications.<\/p>\n<h3 class=\"import-Normal\"><strong>Types of Traits<\/strong><\/h3>\n<p class=\"import-Normal\">When evaluating relationships between taxa, we use key traits that allow us to determine which species are most closely related to one another. Traits can be either ancestral or derived. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1124\">Ancestral traits<\/a><\/strong> are those that a taxon has because it has inherited the trait from a distant ancestor. For example, all primates have body hair because we are mammals and all mammals share an ancestor hundreds of millions of years ago that had body hair. This trait has been passed down to all mammals from a shared ancestor, so all mammals alive today have body hair. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1126\">Derived traits<\/a><\/strong> are those that have been more recently altered. This type of trait is most useful when we are trying to distinguish one group from another because derived traits tell us which taxa are more closely related to each other. For example, humans walk on two legs.The many adaptations that humans possess that allow us to move in this way evolved after humans split from the Genus <em>Pan<\/em>. This means that when we find fossil taxa that share derived traits for walking on two legs, we can conclude that they are likely more closely related to humans than to chimpanzees and bonobos<em>. <\/em><\/p>\n<p class=\"import-Normal\">There are a couple of other important points about ancestral and derived traits that will become apparent as we discuss primate diversity. First, the terms <em>ancestral<\/em> and <em>derived<\/em> are relative terms, meaning that a trait can be either one depending on the taxa being compared. For example, in the previous paragraph, body hair was used as an example for an ancestral trait among primates. All mammals have body hair because we share a distant ancestor who had this trait. The presence of body hair therefore doesn\u2019t allow you to distinguish whether monkeys are more closely related to apes or lemurs because they all share this trait. However, if we are comparing mammals to birds and fish, then body hair becomes a derived trait of mammals. It evolved after mammals diverged from birds and fish, and it tells us that all mammals are more closely related to each other than they are to birds or fish.The second important point is that very often when one lineage splits into two, one taxon will stay more similar to the last common ancestor in retaining more ancestral traits, whereas the other lineage will usually become more different from the last common ancestor by developing more derived traits. This will become very apparent when we discuss the two suborders of primates, Strepsirrhini and Haplorrhini. When these two lineages diverged, strepsirrhines retained more ancestral traits (those present in the earliest primates) and haplorrhines developed more derived traits (became more different from ancestral primates).<\/p>\n<p class=\"import-Normal\">There are two other types of traits that will be relevant to our discussions here: generalized and specialized traits. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1128\">Generalized traits<\/a> <\/strong>are those characteristics that are useful for a wide range of things. Having <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1138\">opposable thumbs<\/a><\/strong> that go in a different direction than the rest of your fingers is a very useful, generalized trait. You can hold a pen, grab a branch, peel a banana, or text your friends all thanks to your opposable thumbs! <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1130\">Specialized traits<\/a> <\/strong>are those that have been modified for a specific purpose. These traits may not have a wide range of uses, but they will be very efficient at their job. Hooves in horses are a good example of a specialized trait: they allow horses to run quickly on the ground on all fours. You can think of generalized traits as a Swiss Army knife, useful for a wide range of tasks but not particularly good at any one of them. That is, if you\u2019re in a bind, then a Swiss Army knife can be very useful to cut a rope or fix a loose screw, but if you were going to build furniture or fix a kitchen sink, then you\u2019d want specialized tools for the job. As we will see, most primate traits tend to be generalized.<\/p>\n<h2 class=\"import-Normal\">What Makes Something a Primate?<\/h2>\n<p class=\"import-Normal\">The Order Primates is distinguished from other groups of mammals in having a <em>suite of characteristics<\/em>. This means that there is no individual trait that you can use to instantly identify an animal as a primate; instead, you have to look for animals that possess a collection of traits. What this also means is that each individual trait we discuss may be found in nonprimates, but if you see an animal that has most or all of these traits, there is a good chance it is a primate.<\/p>\n<p class=\"import-Normal\">Primates are most distinguishable from other organisms in traits related to our vision. Our Order relies on vision as a primary sense, which is reflected in many areas of our anatomy and behaviour. All primates have eyes that face forward with convergent (overlapping) visual fields. So if you cover one eye with your hand, you can still see most of the room with your other one. This also means that we cannot see on the sides or behind us as well as some other animals can. In order to protect the sides of the eyes from the muscles we use for chewing, all primates have at least a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1132\">postorbital bar<\/a>, <\/strong>a bony ring around the outside of the eye (Figure 6.2). Primate taxa with more convergent eyes need extra protection, so animals with greater orbital convergence will have a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1134\">postorbital plate<\/a> <\/strong>or<strong> <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1136\">postorbital closure<\/a> <\/strong>in addition to the bar (Figure 6.2).The postorbital bar is a derived trait of primates, appearing in our earliest ancestors.<\/p>\n<figure style=\"width: 661px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image2-4.jpg\" alt=\"Skulls of a monkey and lemur viewed from the side and top.\" width=\"661\" height=\"430\" \/><figcaption class=\"wp-caption-text\">Figure 6.2: All primates have bony protection around their eyes. Some have a postorbital bar only (right), but many have full postorbital closure, also called a postorbital plate, that completely protects the back of the eye socket (left). Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-4\/\">Postorbital bar\/Postorbital closure (Figure 5.1)<\/a> a derivative work original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Stephanie Etting is 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:\/\/animaldiversity.org\/accounts\/Otolemur_crassicaudatus\/specimens\/collections\/contributors\/phil_myers\/ADW_mammals\/specimens\/Primates\/Otolemur_crassicaudatus\/lateral\/\">Otolemur crassicaudatus (greater galago)<\/a> by <a href=\"https:\/\/animaldiversity.org\/\">Animal Diversity Web<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/3.0\/\">CC BY-NC-SA 3.0<\/a>; <a href=\"https:\/\/animaldiversity.org\/accounts\/Otolemur_crassicaudatus\/specimens\/collections\/contributors\/phil_myers\/ADW_mammals\/specimens\/Primates\/Otolemur_crassicaudatus\/dorsal1809\/\">Macaca fascicularis (long-tailed macaque)<\/a> by <a href=\"https:\/\/animaldiversity.org\/\">Animal Diversity Web<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/3.0\/\">CC BY-NC-SA 3.0<\/a>.]<\/figcaption><\/figure>\n<p class=\"import-Normal\">Another distinctive trait of our Order is that many primates have <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1140\">trichromatic color vision<\/a><\/strong>, the ability to distinguish reds and yellows in addition to blues and greens. Birds, fish, and reptiles are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1142\">tetrachromatic<\/a> <\/strong>(they can see reds, yellows, blues, greens, and even ultraviolet), but most mammals, including some primates, are only <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1144\">dichromatic<\/a> <\/strong>(they see only in blues and greens). It is thought that the nocturnal ancestors of mammals benefited from seeing better at night rather than in colour, and so dichromacy is the ancestral condition for mammals. Trichromatic primates are known to use their colour vision for all sorts of purposes: finding young leaves and ripe fruits, identifying other species, and evaluating signals of health and fertility.<\/p>\n<p class=\"import-Normal\">The primate visual system uses a lot of energy, so primates have compensated by cutting back on other sensory systems, particularly our sense of smell. Compared to other mammals, primates have reduced snouts, another derived trait that appears even in the earliest primate ancestors. There is variation across primate taxa in how much snouts are reduced. Those with a better sense of smell usually have poorer vision than those with a relatively dull sense of smell. The reason for this is that all organisms have a limited amount of energy to spend on running our bodies, so we make <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1146\">evolutionary trade-offs<\/a><\/strong>, as energy spent on one trait cuts back on energy spent on another. So primates with better vision are spending more energy on vision and thus have a poorer smell (and shorter snout), and those who spend less energy on vision will have a better sense of smell (and a longer snout).<\/p>\n<p class=\"import-Normal\">Primates also differ from other mammals in the size and complexity of our brains. On average, primates have brains that are twice as big for their body size when compared to other mammals. Not unexpectedly, the visual centres of the brain are larger in primates and the wiring is different from that in other animals, reflecting our reliance on this sense. The neocortex, which is used for higher functions like consciousness and language in humans, as well as sensory perception and spatial awareness, is also larger in primates relative to other animals. In nonprimates this part of the brain is often smooth, but in primates it is made up of many folds, which increase the surface area. It has been proposed that the more complex neocortex of primates is related to diet, with fruit-eating primates having larger relative brain sizes than leaf-eating primates, due to the more challenging cognitive demands required to find and process fruits (Clutton-Brock &amp; Harvey 1980). An alternative hypothesis argues that larger brain size is necessary for navigating the complexities of primate social life, with larger brains occurring in species who live in bigger, more complex groups relative to those living in pairs or solitarily (Dunbar 1998). There seems to be support for both hypotheses, as large brains are a benefit under both sets of selective pressures.<\/p>\n<p class=\"import-Normal\">Animals with large brains usually have extended life history patterns, and primates are no exception. <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1148\">Life history<\/a> <\/strong>refers to the pace at which an organism grows, reproduces, and ages. Some animals grow very quickly and reproduce many offspring in a short time frame but do not live very long. Other animals grow slowly, reproduce few offspring, reproduce infrequently, and live a long time. Primates are all in the \u201cslow lane\u201d of life history patterns. Compared to animals of similar body size, primates grow and develop more slowly, have fewer offspring per pregnancy, reproduce less often, and live longer. Primates also invest heavily in each offspring. With a few exceptions, most primates only have one offspring at a time. A group of small-bodied monkeys in South America regularly give birth to twins, and some lemurs can give birth to multiple offspring at a time, but these primates are the exception rather than the rule. Primates also reproduce relatively infrequently. The fastest-reproducing primates will produce offspring about every six months, while the slowest, the orangutan, reproduces only once every seven to nine years. This very slow reproductive rate makes the orangutan the slowest-reproducing animal on the planet! Primates are also characterized by having long lifespans. The group that includes humans and large-bodied apes has the most extended life history patterns among all primates, with some large-bodied apes estimated to live up to 58 years in the wild (Robson Et al. 2006).<\/p>\n<figure style=\"width: 392px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image3-5.jpg\" alt=\"Various hands and feet of different primate species.\" width=\"392\" height=\"624\" \/><figcaption class=\"wp-caption-text\">Figure 6.3: These drawings of the hands and feet of different primates show the opposable thumbs and big toes, pentadactyly, flattened nails, and tactile pads characteristic of our Order. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:PrimateFeet.jpg\">PrimateFeet<\/a> by <a href=\"https:\/\/en.wikipedia.org\/wiki\/Richard_Lydekker\">Richard Lydekker<\/a>, original from The Royal Natural History 1:15 (1893), is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p>Primates also differ from other animals in our hands and feet. The Order Primates is a largely <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1150\">arboreal<\/a><\/strong> taxonomic group, meaning that most primates spend a significant amount of their time in trees. As a result, the hands and feet of primates have evolved to move in a three-dimensional environment. Primates have the generalized trait of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1152\">pentadactyly<\/a><\/strong>\u2014 possessing five digits (fingers and toes) on each limb. Many nonprimates, like dogs and horses, have fewer digits because they are specialized for high-speed, <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1154\">terrestrial<\/a><\/strong> (on the ground) running. Pentadactyly is also an ancestral trait, one that dates back to the earliest four-footed animals. Primates today have opposable thumbs and, with the exception of humans, opposable big toes (Figure 6.3). Opposable thumbs and toes are a derived trait that appeared in the earliest primate fossils about 55 million years ago. Having thumbs and big toes that go in a different direction from the rest of the fingers and toes allow primates to be excellent climbers in trees as well as to manipulate objects. Our ability to manipulate objects is further enhanced by the flattened nails on the backs of our fingers and toes that we possess in the place of the claws and hooves that many other mammals have. On the other side of our digits, we have sensitive <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1156\">tactile pads<\/a> <\/strong>that allow us to have a fine sense of touch. Primates use this fine sense of touch for handling food and, in many species, grooming themselves and others. In primates, grooming is an important social currency, through which individuals forge and maintain social bonds.<\/p>\n<table class=\"alignright\" style=\"width: 219.75pt;height: 407px\">\n<caption>Figure 6.4: Primate Traits at a Glance: This table summarizes the suite of traits that differentiate primates from other mammals. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-4\/\">Primate at a glance table (Figure 5.3)<\/a> by Stephanie Etting 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: 30px\">\n<td class=\"a-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 30px;width: 361.667px\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Primate suite of traits<\/strong><\/p>\n<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"a-R\" style=\"height: 362px\">\n<td class=\"a-C\" style=\"background-color: transparent;padding: 5pt;border: 1pt solid #000000;height: 362px;width: 361.667px\">\n<p class=\"import-Normal\" style=\"text-align: center\">Convergent eyes<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Postorbital bar<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Many have trichromatic colour vision<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Short snouts<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Opposable thumbs and big toes<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Pentadactyly<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Flattened nails<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Tactile pads<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Highly arboreal<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Large brains<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Extended life histories<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Live in the tropics<\/p>\n<\/td>\n<\/tr>\n<tr style=\"height: 15px\">\n<td style=\"height: 15px;width: 362.033px\"><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<p>Lastly, primates are very social animals. All primates, even those that search for food alone, establish strong social networks within species. Unlike many animals, primates do not migrate: they stay in a relatively stable area for their whole life, often interacting with the same individuals for their long lives. The long-term relationships that primates form with others of their species lead to complex and fascinating social behaviours (see Chapter 7). Finally, nonhuman primates show a clear preference for tropical regions of the world. Most primates are found between the Tropic of Cancer and the Tropic of Capricorn, with only a few taxa living outside these regions. Figure 6.4 shows a summary of primate traits.<\/p>\n<h2 class=\"import-Normal\">Key Traits Used to Distinguish Between Primate Taxa<\/h2>\n<p class=\"import-Normal\">When placing primate species into specific taxonomic groups, we focus on dental characteristics, behavioural adaptations, and locomotor adaptations. Differences in these characteristics across groups reflect constraints of evolutionary history as well as variation in adaptations.<\/p>\n<h3 class=\"import-Normal\"><strong>Dental Characteristics<\/strong><\/h3>\n<p class=\"import-Normal\">Teeth may not seem like the most exciting topic with which to start, but we can learn a tremendous amount about an organism from its teeth. First, teeth are vital to survival. Wild animals do not have the benefit of knives and forks; they rely on their teeth to process their food. Because of this, teeth of any species have evolved to reflect what that organism eats and therefore have a lot to tell us about their diet. Second, variation in tooth size, shape, and number reveals an organism\u2019s evolutionary history. Some taxa have more teeth than others or different forms of teeth. Furthermore, differences in teeth between males and females can tell us about competition over mates (see Chapter 7). Lastly, teeth are overly represented in the fossil record. Enamel is hard, and there is little meat on jaws so carnivores and scavengers often leave them behind. Sometimes, the only remains we have from an extinct taxon is its teeth!<\/p>\n<figure style=\"width: 356px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image4-4-1.jpg\" alt=\"Yawning baboon with large teeth.\" width=\"356\" height=\"266\" \/><figcaption class=\"wp-caption-text\">Figure 6.5: This picture of an open-mouthed Hamadryas baboon demonstrates the diastema between his upper canine and front teeth. This space is taken up by his lower canine when he closes his mouth. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Ha,ha,ha_...._(14986571843).jpg\">Ha,ha,ha .... (14986571843)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/104249543@N07\">Rolf Dietrich Brecher<\/a> from Germany 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>Like other mammals, primates are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1158\">heterodont<\/a><\/strong>: they have multiple types of teeth that are used for different purposes. We have <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1160\">incisors<\/a> <\/strong>for slicing; <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1162\">premolars<\/a> <\/strong>and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1164\">molars<\/a> <\/strong>for grinding up our food; and <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1166\">canines<\/a><\/strong>, which most primates (not humans) use as weapons against predators and each other. The sizes of canines vary across species and can often be <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1168\">sexually dimorphic<\/a><\/strong>, with males tending to have larger canines than females. Some nonhuman primates <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1170\">hone<\/a><\/strong>, or sharpen, their canines by gnashing the teeth together to sharpen the sides. The upper canine sharpens on the first lower premolar and the lower canine sharpens on the front of the upper canine. As canines get larger, they require a space to fit in order for the jaws to close. This space between the teeth is called a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1172\">diastema<\/a><\/strong> (Figure 6.5).<\/p>\n<p class=\"import-Normal\">We use a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1174\">dental formula<\/a><\/strong> to specify how many incisors, canines, premolars, and molars are in each quadrant of the mouth (half of the top or bottom). For example, Figure 6.6 shows half of the lower teeth of a human. You can see that in half of the mandible, there are two incisors, one canine, two premolars, and three molars. This dental formula is written as 2:1:2:3. (The first number represents the number of incisors, followed by the number of canines, premolars, and molars.)<\/p>\n<figure style=\"width: 241px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image5-5.png\" alt=\"Human mandible with four types of teeth.\" width=\"241\" height=\"424\" \/><figcaption class=\"wp-caption-text\">Figure 6.6:\u00a0 This drawing shows half of the human mandible. With the four types of teeth labeled, you can determine that the dental formula is 2:1:2:3. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Gray997.png\">Gray997<\/a> by <a href=\"https:\/\/en.wikipedia.org\/wiki\/Henry_Vandyke_Carter\">Henry Vandyke Carter<\/a>, original in Henry Gray (1918) Anatomy of the Human Body, Plate 997, is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">To determine the dental formula, you need to be able to identify the different types of teeth. You can recognize incisors because they often look like spatulas with a flat, blade-like surface. Premolars and molars can be differentiated by the number of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1176\">cusps<\/a><\/strong> that they have. Cusps are the bumps that you can feel with your tongue on the surface of your back teeth. Premolars are smaller than molars and, in primates, often have one or two cusps on them. Molars are bigger, providing a larger chewing surface, and have more cusps. Depending on the species and whether you\u2019re looking at upper or lower teeth, primate molars can have between three and five cusps. Molar cusps can also vary between taxa in how they are arranged; you will learn more about this later in this chapter. Canines are often easy to distinguish because, in most taxa, they are much longer and more conical than the other teeth.<\/p>\n<p class=\"import-Normal\">Teeth also directly reflect an organism\u2019s diet. Primates are known to eat a wide range of plant parts, insects, gums, and, rarely, meat. While all primates eat a variety of foods, what differs among primates are the proportions of each of these food items in the diet. That is, two primates living in the same forest may be eating the same foods but in vastly different proportions, and so we would categorize them as different dietary types. The most common dietary types among primates are those whose diets consist primarily of fruit (<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_954\">frugivores<\/a><\/strong>), those who eat mostly insects (<strong>insectivores<\/strong>), and those who eat primarily leaves (<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_956\">folivores<\/a><\/strong>). A few primate taxa are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1178\">gummivores<\/a><\/strong>, specializing in eating gums and saps, but we will only focus on the adaptations found in the three primary dietary groups.<\/p>\n<h4 class=\"import-Normal\"><em>Frugivores<\/em><\/h4>\n<p class=\"import-Normal\">Plants want animals to eat their fruits because, in doing so, animals eat the seeds of the fruit and then disperse them far away from the parent plant. Therefore, plants often \u201cadvertise\u201d fruits by making them colourful and easy to spot, full of easy-to-digest sugars that make them taste good and, often, easy to chew and digest (not being too fibrous or tough). For these reasons, frugivores often do not need a lot of specialized traits to consume a diet rich in fruits (Figure 6.7). Their molars usually have a broad chewing surface with low, rounded cusps (referred to as <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1180\">bunodont<\/a> <\/strong>molars). Frugivores have large incisors for slicing through the outer coatings on fruit, and they tend to have stomachs, colons, and small intestines that are intermediate in terms of size and complexity between insectivores and folivores (Chivers &amp; Hladik 1980). They are also usually of intermediate body size between the other two dietary types. Because fruit does not contain protein, frugivores must supplement their diet with protein from insects, leaves, and\/or seeds. Frugivores who get protein by eating seeds evolved to have thicker enamel on their teeth to protect them from excessive wear.<\/p>\n<figure style=\"width: 503px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image6-2.jpg\" alt=\"Upper teeth and maxilla of a frugivore monkey.\" width=\"503\" height=\"331\" \/><figcaption class=\"wp-caption-text\">Figure 6.7: Frugivores are characterized by large incisors, bunodont molars, and digestive tracts that are intermediate in complexity between the other two dietary types. Credit: <a href=\"https:\/\/animaldiversity.org\/collections\/contributors\/phil_myers\/ADW_mammals\/specimens\/Primates\/Papio_papio\/utr0087\/\">Papio papio (Guinea baboon).jpg<\/a> by Phil Myers on <a href=\"https:\/\/animaldiversity.org\">Animal Diversity Web<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/3.0\/\">CC BY-NC-SA 3.0 License. <\/a><\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Insectivores<\/em><\/h4>\n<p class=\"import-Normal\">While insects can be difficult to find and catch, they are easy to chew and digest. As a result, insectivorous primates usually have small molars with pointed cusps to puncture the exoskeleton of the insects (Figure 6.8), and they have simple stomachs and colons with a long small intestine to process the insects. Nutritionally, insects provide a lot of protein and fat but are not plentiful enough in the environment to support large-bodied animals, so insectivores are usually the smallest of the primates.<\/p>\n<figure style=\"width: 351px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image7-3.jpg\" alt=\"Mandible, upper teeth, and maxilla of insectivore tarsier.\" width=\"351\" height=\"280\" \/><figcaption class=\"wp-caption-text\">Figure 6.8: Insectivores need sharp, pointed molar cusps to break through the exoskeletons of insects. Insects are easy to digest, so these primates have simple digestive tracts. Credit: Tarsier (an insectivor)\u2019s teeth original to Explorations: An Open Invitation to Biological Anthropology (2nd ed.) by Stephanie Etting is a collective work under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/3.0\/\">CC BY-NC-SA 3.0 License.<\/a> [Includes <a href=\"https:\/\/animaldiversity.org\/collections\/contributors\/phil_myers\/ADW_mammals\/specimens\/Primates\/Tarsius_syrichta\/lower_lateral1942\/\">Lower_lateral1942<\/a> by Phil Myers on <a href=\"https:\/\/animaldiversity.org\">Animal Diversity Web<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/3.0\/\">CC BY-NC-SA 3.0<\/a>; <a href=\"https:\/\/animaldiversity.org\/collections\/contributors\/phil_myers\/ADW_mammals\/specimens\/Primates\/Tarsius_syrichta\/ventral\/\">Ventral<\/a> by Phil Myers on <a href=\"https:\/\/animaldiversity.org\">Animal Diversity Web<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/3.0\/\">CC BY-NC-SA 3.0<\/a>.]<\/figcaption><\/figure>\n<h4 class=\"import-Normal\"><em>Folivores<\/em><\/h4>\n<p class=\"import-Normal\">Plants rely on leaves to get energy from the sun, so plants do not want animals to eat their leaves (unlike their fruit). As a result, plants evolved to try to discourage animals from eating their leaves. Leaves often carry toxins, taste bitter, are very fibrous and difficult to chew, and are made of large cellulose molecules that are difficult to break down into usable sugars. Thus, animals who eat leaves need a lot of specialized traits (Figure 6.9). Folivorous primates have broad molars with high, sharp cusps connected by <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1182\">shearing crests<\/a><\/strong>. These molar traits allow folivores to physically break down fibrous leaves when chewing. Folivores then chemically break down cellulose molecules into usable energy. To do this, some folivores have complex stomachs with multiple compartments, while others have large, long intestines and special gut bacteria that can break up cellulose. Folivores are usually the largest bodied of all primates, and they tend to spend a large portion of their day digesting their food, so they are less active than frugivores or insectivores.<\/p>\n<figure style=\"width: 468px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image8-2.jpg\" alt=\"Upper teeth and maxilla of a monkey shows folivore traits.\" width=\"468\" height=\"337\" \/><figcaption class=\"wp-caption-text\">Figure 6.9: To derive energy from leaves, folivores, like this Trachypithecus (dusky leaf monkey), have smaller incisors and high sharp molar cusps connected by shearing crests. Credit: <a href=\"https:\/\/animaldiversity.org\/collections\/contributors\/phil_myers\/ADW_mammals\/specimens\/Primates\/Trachypithecus_obscurus\/utr0075\/\">Trachypithecus obscurus (dusky leaf monkey) upper teeth<\/a> by Phil Myers on <a href=\"https:\/\/animaldiversity.org\">Animal Diversity Web<\/a> has been modified (background removed, labels added by Stephanie Etting) is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/3.0\/\">CC BY-NC-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Behavioural Adaptations<\/strong><\/h3>\n<p class=\"import-Normal\">Since Chapter 6 is dedicated to primate behaviour, we will only briefly discuss variations in activity patterns, social grouping, and habitat use. Primate groups differ in <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1184\">activity patterns<\/a><\/strong>: whether they are active during the day (<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1012\">diurnal<\/a><\/strong>), at night (<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1010\">nocturnal<\/a><\/strong>), or through the 24-hour period (<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1186\">cathemeral<\/a><\/strong>). Primate taxa vary in social groupings: some are primarily solitary, others live in pairs, and still others live in groups of varying sizes and compositions. Lastly, some taxa are primarily arboreal while others are more terrestrial.<\/p>\n<h3 class=\"import-Normal\"><strong>Locomotor Adaptations<\/strong><\/h3>\n<p class=\"import-Normal\">Finally, primate groups vary in their adaptations for different forms of <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1188\">locomotion<\/a><\/strong>, or how they move around. Living primates are known to move by vertical clinging and leaping, quadrupedalism, brachiation, and bipedalism.<\/p>\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1190\">Vertical clinging and leaping<\/a> <\/strong>is when an animal grasps a vertical branch with its body upright, pushes off with long hind legs, and then lands on another vertical support branch (Figure 6.10a). Animals who move in this way usually have longer legs than arms, long fingers and toes, and smaller bodies. Vertical clinger leapers also tend to have elongated ankle bones, which serve as a lever to help them push off with their legs and leap to another branch (Figure 6.10b).<\/p>\n<\/div>\n<figure id=\"attachment_181\" aria-describedby=\"caption-attachment-181\" style=\"width: 608px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-147\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/5.10.jpg\" alt=\"Movement of vertical clinger and leaper, and tarsier skeleton.\" width=\"608\" height=\"462\" \/><figcaption id=\"caption-attachment-181\" class=\"wp-caption-text\">Figure 6.10a: Vertical clingers and leapers have longer legs than arms, long lower backs, and long fingers and toes. They also have elongated ankle bones to help them push off when leaping. Credit: a. <a href=\"https:\/\/upload.wikimedia.org\/wikipedia\/commons\/6\/6e\/Propithecus_vertical_clinging_and_leaping.svg\">Propithecus vertical clinging and leaping<\/a> by Terpsichores is under a<a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/\"> CC BY-SA 3.0 License<\/a>. 6.10b: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Em_-_Tarsius_tarsier_-_3.jpg\">Tarsier skeleton<\/a> by Em\u0151ke D\u00e9nes has been modified (background removed) by Stephanie Etting and is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/\">CC BY-SA 4.0 License<\/a>. Original Spectral tarsier (Tarsius tarsier) skeleton at the Cambridge University Museum of Zoology, England.)<\/figcaption><\/figure>\n<p>&nbsp;<\/p>\n<div class=\"learning-objectives\">\n<p class=\"import-Normal\"><strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1192\">Quadrupedalism<\/a><\/strong>, walking on all fours, is the most common form of locomotion among primates. Quadrupedal animals usually have legs and arms that are about the same length and a tail for balance. Arboreal quadrupeds (Figure 6.11a) usually have shorter arms and legs and longer tails, while terrestrial quadrupeds (Figure 6.11b) have longer arms and legs and, often, shorter tails. These differences relate to the lower centre of gravity needed by arboreal quadrupeds for balance in trees and the longer tail required for better balance when moving along the tops of branches. Terrestrial quadrupeds have longer limbs to help them cover more distance more efficiently.<\/p>\n<\/div>\n<figure id=\"attachment_181-2\" aria-describedby=\"caption-attachment-181-2\" style=\"width: 704px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-148\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/5.11.jpg\" alt=\"Arboreal quadrupedal monkey and terrestrial quadrupedal monkey.\" width=\"704\" height=\"251\" \/><figcaption id=\"caption-attachment-181-2\" class=\"wp-caption-text\">Figure 6.11a\/b: Two examples of quadrupedal primates. The capuchin monkey skeleton on the left (a) is a typical arboreal quadruped with shorter arms and legs, longer fingers and toes, and a long tail. The baboon skeleton on the right (b) is a terrestrial quadruped with relatively long arms and legs, shorter fingers and toes, and a short tail. Credits: 6.11a. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Description_iconographique_compar%C3%A9e_du_squelette_et_du_syst%C3%A8me_dentaire_des_mammif%C3%A8res_r%C3%A9cents_et_fossiles_(Sapajus_apella).jpg\">Capuchin monkey skeleton<\/a> by Henri-Marie Ducrotay de Blainville is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>. 6.11b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Description_iconographique_compar%C3%A9e_du_squelette_et_du_syst%C3%A8me_dentaire_des_mammif%C3%A8res_r%C3%A9cents_et_fossiles_(Papio_ursinus).jpg\">Baboon<\/a> by Henri-Marie Ducrotay de Blainville is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<div class=\"learning-objectives\">\n<p class=\"import-Normal\">The third form of locomotion seen in primates is<strong> <a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1194\">brachiation<\/a><\/strong>, the way of moving you used if you played on \u201cmonkey bars\u201d as a child. Brachiation involves swinging below branches by the hands (Figure 6.12a). To be an efficient brachiator, a primate needs to have longer arms than legs, flexible shoulders and wrists, a short lower back, and no tail (Figure 6.12b). Some primates move via <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1196\">semi-brachiation<\/a><\/strong>, in which they swing below branches but do not have all of the same specializations as brachiators. Semi-brachiators have flexible shoulders, but their arms and legs are about the same length, which is useful because they are quadrupedal when on the ground. They also use long <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1198\">prehensile tails<\/a><\/strong> as a third limb when swinging (Figure 6.13). The underside of the tail has a tactile pad, resembling your fingerprints, for better grip.<\/p>\n<\/div>\n<figure id=\"attachment_181-3\" aria-describedby=\"caption-attachment-181-3\" style=\"width: 1600px\" class=\"wp-caption alignnone\"><img class=\"wp-image-149 size-full\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/5.12.jpg\" alt=\"Primate swinging through branches and gibbon skeleton.\" width=\"1600\" height=\"800\" \/><figcaption id=\"caption-attachment-181-3\" class=\"wp-caption-text\">Figure 6.12a. Example of brachiation. 6.12b. Skeleton of a typical brachiator, showing longer arms than legs, short back, and lack of a tail. Credit: a. <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-4\/\">Brachiator (Figure 5.9b)<\/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>. b. <a href=\"https:\/\/upload.wikimedia.org\/wikipedia\/commons\/a\/a3\/Giboia.jpg\">Skeleton of Gibbon (Giboia) <\/a>by Joxerra Aihartza is under a <a href=\"https:\/\/artlibre.org\">Free Art License<\/a>.<\/figcaption><\/figure>\n<div class=\"learning-objectives\">\n<figure style=\"width: 565px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image15-1.jpg\" alt=\"Spider monkey swinging below a rope.\" width=\"565\" height=\"377\" \/><figcaption class=\"wp-caption-text\">Figure 6.13. Spider monkeys are considered semi-brachiators, as they can swing below branches but use their tails as a third limb. On the ground they move via quadrupedal locomotion. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Ateles-fusciceps_54724770b.jpg\">Ateles-fusciceps 54724770b<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:LeaMaimone\">LeaMaimone<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.5\/legalcode\">CC BY 2.5 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Lastly, humans move around on two feet, called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1200\">bipedalism<\/a><\/strong>. Some nonhuman primates will occasionally travel on two feet but do so awkwardly and never for long distances. Among mammals, only humans have evolved to walk with a striding gait on two legs as a primary form of locomotion.<\/p>\n<h2 class=\"import-Normal\">Primate Diversity<\/h2>\n<p class=\"import-Normal\">As we begin exploring the different taxa of primates, it is important to keep in mind the hierarchical nature of taxonomic classification and how this relates to the key characteristics that will be covered. Figure 6.14 summarizes the major taxonomic groups of primates that you will learn about here. If you locate humans on the chart, you can trace our classification and see all of the categories getting more inclusive as you work your way up to the Order Primates. This means that humans will have the key traits of each of those groups. It is a good idea to refer to the figure to orient yourself as we discuss each taxon.<\/p>\n<figure style=\"width: 2048px\" class=\"wp-caption alignnone\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image16-1.jpg\" alt=\"Taxonomic chart shows primate order, suborder, infraorder, superfamily, and species.\" width=\"2048\" height=\"1154\" \/><figcaption class=\"wp-caption-text\">Figure 6.14: This taxonomy chart shows the major groups of primate taxa, starting with the largest category (Order) and moving to more specific categories and examples. <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__-4\/\">Primate taxonomy char (Figure 5.11)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Stephanie Etting is a collective work under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/\">CC BY-NC 4.0 License<\/a>. [Includes <a href=\"https:\/\/phylopic.org\/image\/d6cfb28f-136e-4a20-a5ac-8eb353c7fc4a\/\">Lemur catta Linnaeus, 1759<\/a> by Roberto D\u00edaz Sibaja, <a href=\"https:\/\/creativecommons.org\/licenses\/by\/3.0\/\">CC BY 3.0<\/a>; <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-4\/\">Lorisoidea<\/a> original to<a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\"> Explorations: An Open Invitation to Biological Anthropology<\/a> by Katie Nelson, <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0<\/a>; <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-4\/\">Tarsiiformes<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Mary Nelson, <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0<\/a>; <a href=\"https:\/\/phylopic.org\/image\/156b515d-f25c-4497-b15b-5afb832cc70c\/\">Cebinae Bonaparte, 1831<\/a> by Sarah Werning, <a href=\"https:\/\/creativecommons.org\/licenses\/by\/3.0\/\">CC BY 3.0<\/a>; <a href=\"https:\/\/phylopic.org\/image\/899742c2-9a40-4fa0-b2cd-2eb133c8f6e8\/\">Colobus guereza Ruppell, 1835<\/a> by Yan Wong, designated to the <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/\">public domain (CC0)<\/a>; <a href=\"https:\/\/phylopic.org\/image\/72f2f854-f3cd-4666-887c-35d5c256ab0f\/\">Papio cynocephalus<\/a> by Owen Jones, designated to the <a href=\"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/\">public domain (CC0)<\/a>; <a href=\"https:\/\/pixabay.com\/vectors\/animals-silhouette-wolf-elephant-2755766\/\">animals silhouette wolf elephant (2755766)<\/a> by <a href=\"https:\/\/pixabay.com\/users\/mohamed_hassan-5229782\/\">mohamed_hassan<\/a>, <a href=\"https:\/\/pixabay.com\/service\/terms\/#license\">Pixabay License<\/a>.]<\/figcaption><\/figure>\n<h3 class=\"import-Normal\"><strong>Suborder Strepsirrhini<\/strong><\/h3>\n<figure style=\"width: 387px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image17-1.jpg\" alt=\"Eight strepsirrhine species.\" width=\"387\" height=\"605\" \/><figcaption class=\"wp-caption-text\">Figure 6.15: (Clockwise from top right) sifaka, black-and-white ruffed lemur, loris, galago, slender loris, mouse lemur, aye-aye, and ring-tailed lemur. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Extant_Strepsirrhini.jpg\">Extant Strepsirrhini<\/a> a collective work by <a href=\"https:\/\/www.flickr.com\/people\/23661161@N02\">Mark Dumont<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>. [Includes <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Katta_csal%C3%A1d.jpg\">Katta csal\u00e1d<\/a> by Veszpr\u00e9mi \u00c1llatkert, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Aye-aye_at_night_in_the_wild_in_Madagascar.jpg\">Aye-aye at night in the wild in Madagascar<\/a> by Frank Vassen, <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/deed.en\">CC BY 2.0<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Diademed_ready_to_push_off.jpg\">Diademed ready to push off<\/a> by Michael Hogan, designated to the <a href=\"https:\/\/creativecommons.org\/share-your-work\/public-domain\/cc0\/\">public domain (CC0)<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Juvenile_Black-and-White_Ruffed_Lemur,_Mantadia,_Madagascar.jpg\">Juvenile Black-and-White Ruffed Lemur, Mantadia, Madagascar<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/42244964@N03\">Frank Vassen<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\">CC BY 2.0<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Microcebus_murinus_-Artis_Zoo,_Amsterdam,_Netherlands-8a.jpg\">Microcebus murinus -Artis Zoo, Amsterdam, Netherlands-8a<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/46956042@N00\">Arjan Haverkamp<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\">CC BY 2.0<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Slow_Loris.jpg\">Slow Loris<\/a> by Jmiksanek, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Slender_Loris.jpg\">Slender Loris<\/a> by Kalyan Varma (<a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Kalyanvarma\">Kalyanvarma<\/a>), <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Garnett's_Galago_(Greater_Bushbaby).jpg\">Garnett's Galago (Greater Bushbaby)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/23661161@N02\">Mark Dumont<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\">CC BY 2.0<\/a>.]<\/figcaption><\/figure>\n<p>The Order Primates is subdivided into Suborder Strepsirrhini and Suborder Haplorrhini, which, according to molecular estimates, split about 70\u201380 million years ago (Pozzi et al. 2014). The strepsirrhines include the groups commonly called lemurs, lorises, and galagos (Figure 6.15). Strepsirrhines differ from haplorrhines in many ways, most of which involve retaining ancestral traits from the earliest primates. Strepsirrhines do have two key derived traits that evolved after they diverged from the haplorrhines: the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1202\">grooming claw<\/a> <\/strong>(Figure 6.16) on the second digit of each foot, and the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1204\">tooth comb<\/a><\/strong> (or <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1206\">dental comb<\/a><\/strong>) located on the lower, front teeth (Figure 6.17). In most strepsirrhines, there are six teeth in the toothcomb\u2014four incisors and two canines. Other than the tooth comb, the teeth of strepsirrhines are fairly simple and are neither large or distinctive relative to haplorrhines.<\/p>\n<figure style=\"width: 237px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image18.jpg\" alt=\"A long, thin dark claw is visible in contrast to flat dark nails on the other digits.\" width=\"237\" height=\"202\" \/><figcaption class=\"wp-caption-text\">Figure 6.16: The foot of a ring-tailed lemur showing its grooming claw on the second digit. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Lemur_catta_toilet_claw.jpg\">Lemur catta toilet claw<\/a> by Alex Dunkel (Maky) is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/3.0\/legalcode\">CC BY 3.0 License.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\">Compared to haplorrhines, strepsirrhines rely more on nonvisual senses. Strepsirrhines get their name because they have wet noses (<strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1208\">rhinariums<\/a><\/strong>) like cats and dogs, a trait that, along with a longer snout, reflect strepsirrhines\u2019 greater reliance on olfaction relative to haplorrhines. Many strepsirrhines use <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1210\">scent marking<\/a><\/strong>, including rubbing scent glands or urine on objects in the environment to communicate with others. Additionally, many strepsirrhines have mobile ears that they use to locate insect prey and predators. While strepsirrhines have a better sense of smell than haplorrhines, their visual adaptations are more ancestral. Strepsirrhines have less convergent eyes than haplorrhines and therefore all have postorbital bars, whereas haplorrhines have full postorbital closure (see Figure 6.2). All strepsirrhines have a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1212\">tapetum lucidum<\/a><\/strong>, a reflective layer at the back of the eye that reflects light and thereby enhances the ability to see in low-light conditions. It is the same layer that causes your dog or cat to have \u201cyellow eye\u201d when you take photos of them with the flash on. This is a trait thought to be ancestral among mammals as a whole.<\/p>\n<figure style=\"width: 292px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image19-1.jpg\" alt=\"The lower, front teeth are long, thin, tightly together in a line, and project towards the lips.\" width=\"292\" height=\"354\" \/><figcaption class=\"wp-caption-text\">Figure 6.17: The lower front teeth of a ring-tailed lemur showing the six teeth of the tooth comb: four incisors and two canines. The teeth that superficially look like canines are premolars. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Lemur_catta_toothcomb.jpg\">Lemur catta toothcomb<\/a> by Alex Dunkel (Maky) is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/3.0\/legalcode\">CC BY 3.0 license.<\/a><\/figcaption><\/figure>\n<p class=\"import-Normal\">Strepsirrhines also differ from haplorrhines in some aspects of their ecology and behaviour. The majority of strepsirrhines are solitary, traveling alone to search for food; a few taxa are more social. Most strepsirrhines are also nocturnal and arboreal. Strepsirrhines are, on average, smaller than haplorrhines, and so many of them have a diet consisting of insects and fruit, with few taxa eating primarily leaves. Lastly, most strepsirrhines are good at leaping, with several taxa specialized for vertical clinging and leaping. In fact, among primates, all but one of the vertical clinger leapers belong to the Suborder Strepsirrhini.<\/p>\n<p class=\"import-Normal\">Strepsirrhines can be found all across Asia, Africa, and on the island of Madagascar (Figure 6.18). The Suborder Strepsirrhini is divided into two groups: (1) the lemurs of Madagascar and (2) the lorises, pottos, and galagos of Africa and Asia. By molecular estimates, these two groups split about 65 million years ago (Pozzi Et al. 2014).<\/p>\n<figure style=\"width: 443px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image20-1.png\" alt=\"Map strepsirrhine primates locations.\" width=\"443\" height=\"342\" \/><figcaption class=\"wp-caption-text\">Figure 6.18: Geographic distribution of living strepsirrhines. Lemurs live only on Madagascar, while lorises and galagos live across Central Africa and South and Southeast Asia. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-4\/\">Geographic distribution of living strepsirrhines (Figure 5.16)<\/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<h4><em>Lemurs of Madagascar<\/em><\/h4>\n<p class=\"import-Normal\">Madagascar is an island off the east coast of Africa, and it is roughly the size of California, Oregon, and Washington combined. It has been separated from Africa for about 130 million years and from India for about 85 million years, which means it was already an island when strepsirrhines got there approximately 60\u201370 million years ago. Only a few mammal species ever reached Madagascar, and so when lemurs arrived they were able to flourish into a variety of forms.<\/p>\n<p class=\"import-Normal\">The lemurs of Madagascar are much more diverse compared to their mainland counterparts, the lorises and galagos. While many Malagasy strepsirrhines are nocturnal, plenty of others are diurnal or cathemeral. They range in body size from the smallest of all primates, the mouse lemur, some species of which weigh a little over an ounce (see Figure 6.15), up to the largest of all strepsirrhines, the indri, which weighs up to about 20 pounds (Figure 6.19). Lemurs include species that are insectivorous, frugivorous, and folivorous. A couple of members of this group have unusual diets for primates, including the gummivorous fork-marked and bamboo lemurs, who are able to metabolize the cyanide in bamboo. The most unique lemur is the aye-aye (depicted in Figure 6.15). This nocturnal lemur has rodent-like front teeth that grow continuously and a long-bony middle finger that it uses to fish grubs out of wood. It has a very large brain compared to other strepsirrhines, which it fuels with a diet that includes bird\u2019s eggs and other animal matter. Based on genetic estimates and morphological studies, it is believed that aye-ayes were the first lemurs to separate from all other strepsirrhines and to evolve on their own since strepsirrhines arrived in Madagascar (Matsui Et al. 2009).<\/p>\n<figure style=\"width: 326px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image21-1-1.jpg\" alt=\"Two Indis in a tree.\" width=\"326\" height=\"217\" \/><figcaption class=\"wp-caption-text\">Figure 6.19: Indris, the largest of the lemurs. These folivorous lemurs are vertical clingers and leapers and live in pairs. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Indri_indri_0003.jpg\">Indri indri 0003<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Christophe_Germain\">Christophe Germain<\/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\">Lemurs are also diverse in terms of social behaviour: Many lemurs are solitary foragers, some live in pairs, others in small groups, still others in larger groups, and some, like the red-ruffed lemur, live in unique and complex social groups (Vasey 2006). Lemurs include some of the best vertical clingers and leapers, and while many lemurs are quadrupedal, even the quadrupedal lemurs are quite adept at leaping. Malagasy strepsirrhines also exhibit a few unusual traits. They are highly seasonal breeders, often mating only during a short window once a year (Wright 1999). Female ring-tailed lemurs, for example, come into estrus one day a year for a mere six hours. Unlike most primates, where males are typically large and dominant, Malagasy strepsirrhines feature socially dominant females that are similar in size to males and have priority access to resources.<\/p>\n<h4 class=\"import-Normal\"><em>Lorises, Pottos, and Galagos of Asia and Africa<\/em><\/h4>\n<figure style=\"width: 207px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image22-1.jpg\" alt=\"Slow loris hanging from a branch.\" width=\"207\" height=\"309\" \/><figcaption class=\"wp-caption-text\">Figure 6.20: This slow loris, like all others in this taxonomic group, is solitary and nocturnal, with a diet heavy in insects and fruit. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Nycticebus_coucang_002.jpg\">Nycticebus coucang 002<\/a> by David Haring \/ Duke Lemur Center 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>Unlike the lemurs of Madagascar, lorises, pottos, and galagos live in areas where they share their environments with monkeys and apes, who often eat similar foods. Lorises live across South and Southeast Asia, while pottos and galagos live across Central Africa. Because of competition with larger-bodied monkeys and apes, mainland strepsirrhines are more restricted in the niches they can fill in their environments and so are less diverse than the lemurs.<\/p>\n<p>The strepsirrhines of Africa and Asia are all nocturnal and solitary, with little variation in body size and diet. For the most part, the diet of lorises, pottos, and galagos consists of fruits and insects. A couple of species eat more gum, but overall the diet of this group is narrow when compared to the Malagasy lemurs. Lorises (Figure 6.20) and pottos are known for being slow, quadrupedal climbers, moving quietly through the forests to avoid being detected by predators. These strepsirrhines have developed additional defences against predators. Lorises, for example, eat a lot of caterpillars, which makes their saliva slightly toxic. Loris mothers bathe their young in this toxic saliva, making the babies unappealing to predators. In comparison to the slow-moving lorises and pottos, galagos are active quadrupedal runners and leapers that scurry about the forests at night. Galagos make distinctive calls that sound like a baby crying, which has led to their nickname \u201cbushbabies.\u201d Figure 6.21 summarizes the key differences between these two groups of strepsirrhines.<\/p>\n<table class=\"aligncenter\" style=\"width: 468pt\">\n<caption>Figure 6.21: Strepsirrhini at a glance: This table summarizes the key differences between the two groups of strepsirrhines. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-4\/\">Strepsirrhines at a glance table (Figure 5.19)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Stephanie Etting 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:Ringtailed_Lemurs_in_Berenty.jpg\">Ringtailed Lemurs in Berenty<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/50852241@N00\">David Dennis<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/legalcode\">CC BY-SA 2.0<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Komba_u%C5%A1at%C3%A1.jpg\">Komba u\u0161at\u00e1<\/a> by Petr Hamern\u00edk, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0<\/a>.]<\/caption>\n<thead>\n<tr>\n<td class=\"a0-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">\n<\/td>\n<td class=\"a0-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong><img class=\"alignnone\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image23-1.jpg\" alt=\"Baby primate on the back of adult primate.\" width=\"231\" height=\"165\" \/><\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Lemurs<\/strong><\/p>\n<\/td>\n<td class=\"a0-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong><img class=\"alignnone\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image24-1.jpg\" alt=\"Small primate with big eyes and long tail.\" width=\"198\" height=\"132\" \/><\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Lorises, Pottos, and Galagos<\/strong><\/p>\n<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"a0-R\">\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Geographic range<\/strong><\/p>\n<\/td>\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Madagascar<\/p>\n<\/td>\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">South and Southeast Asia<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Central Africa<\/p>\n<\/td>\n<\/tr>\n<tr class=\"a0-R\">\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Activity patterns<\/strong><\/p>\n<\/td>\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Diurnal, nocturnal, or cathemeral<\/p>\n<\/td>\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Nocturnal<\/p>\n<\/td>\n<\/tr>\n<tr class=\"a0-R\">\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Dietary types<\/strong><\/p>\n<\/td>\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Insectivore, frugivore, or folivore<\/p>\n<\/td>\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Insectivore, frugivore<\/p>\n<\/td>\n<\/tr>\n<tr class=\"a0-R\">\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Social groupings<\/strong><\/p>\n<\/td>\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Solitary, pairs, or small to large groups<\/p>\n<\/td>\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Solitary<\/p>\n<\/td>\n<\/tr>\n<tr class=\"a0-R\">\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Forms of locomotion<\/strong><\/p>\n<\/td>\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Vertical clinger leapers, quadrupedal<\/p>\n<\/td>\n<td class=\"a0-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Slow quadrupedal climbers and active quadrupedal runners<\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<h3 class=\"import-Normal\"><strong>Suborder Haplorrhini<\/strong><\/h3>\n<p class=\"import-Normal\">When the two primate suborders split from one another, strepsirrhines retained more ancestral traits while haplorrhines developed more derived traits, which are discussed below.<\/p>\n<p class=\"import-Normal\">As mentioned earlier, haplorrhines have better vision than strepsirrhines. This is demonstrated by the full postorbital closure protecting the more convergent eyes that haplorrhines possess (with one exception seen in Figure 6.2). Most haplorrhines are trichromatic, and all have a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1410\">fovea<\/a><\/strong>, a depression in the retina at the back of the eye containing concentrations of cells that allows them to see things very close up in great detail. The heavier reliance on vision over olfaction is also reflected in the shorter snouts ending with the <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1412\">dry nose<\/a> <\/strong>(no rhinarium) of haplorrhines. All but two genera of living haplorrhines are active during the day, so this group lacks the tapetum lucidum that is so useful to nocturnal species. On average, haplorrhines also have larger brains relative to their body size when compared with strepsirrhines.<\/p>\n<p class=\"import-Normal\">The Haplorrhini differ from the Strepsirrhini in their ecology and behaviour as well. Haplorrhines are generally larger than strepsirrhines, and they tend to be folivorous and frugivorous. This dietary difference is reflected in the teeth of haplorrhines, which are broader with more surface area for chewing. The larger body size of this taxon also influences locomotion. Only one haplorrhine is a vertical clinger and leaper. Most members of this suborder are quadrupedal, with one subgroup specialized for brachiation. A few haplorrhine taxa are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1414\">monomorphic<\/a><\/strong>, meaning males and females are the same size, but many members of this group show moderate to high sexual dimorphism in body size and canine size. Haplorrhines also differ in social behaviour. All but two haplorrhines live in groups, which is very different from the primarily solitary strepsirrhines. Differences between the two suborders are summarized in Figure 6.22.<\/p>\n<table class=\"aligncenter\" style=\"width: 468pt\">\n<caption>Figure 6.22: Suborders at a glance: This table summarizes the key differences between the two primate suborders. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-4\/\">Suborders at a glance table (Figure 5.20)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Stephanie Etting 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:Black-and-White_Ruffed_Lemur,_Mantadia,_Madagascar.jpg\">Black-and-White Ruffed Lemur, Mantadia, Madagascar<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/42244964@N03\">Frank Vassen<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\">CC BY 2.0<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Crab_eating_macaque_face.jpg\">Crab eating macaque face<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Bruce89\">Bruce89<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0<\/a>.]<\/caption>\n<thead>\n<tr>\n<td class=\"a1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">\n<\/td>\n<td class=\"a1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong><img class=\"alignnone\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image25.jpg\" alt=\"Black-and-white ruffed lemur.\" width=\"213\" height=\"159\" \/><\/strong><strong>Suborder Strepsirrhini<\/strong><\/p>\n<\/td>\n<td class=\"a1-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong><img class=\"alignnone\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image26-1.jpg\" alt=\"Crab-eating macaque.\" width=\"137\" height=\"137\" \/><\/strong><\/p>\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Suborder Haplorrhini<\/strong><\/p>\n<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"a1-R\">\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Sensory adaptations<\/strong><\/p>\n<\/td>\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Rhinarium<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Longer snout<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Eyes less convergent<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Postorbital bar<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Tapetum lucidum<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Mobile ears<\/p>\n<\/td>\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">No rhinarium<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Short snout<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Eyes more convergent<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Postorbital plate<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">No tapetum lucidum<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Many are trichromatic<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Fovea<\/p>\n<\/td>\n<\/tr>\n<tr class=\"a1-R\">\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Dietary differences<\/strong><\/p>\n<\/td>\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Mostly insectivores and frugivores, few folivores<\/p>\n<\/td>\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Few insectivores, mostly frugivores and folivores<\/p>\n<\/td>\n<\/tr>\n<tr class=\"a1-R\">\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Activity patterns and Ecology<\/strong><\/p>\n<\/td>\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Mostly nocturnal, few diurnal or cathemeral<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Almost entirely arboreal<\/p>\n<\/td>\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Only two are nocturnal, rest are diurnal<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Many arboreal taxa, also many terrestrial taxa<\/p>\n<\/td>\n<\/tr>\n<tr class=\"a1-R\">\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Social groupings<\/strong><\/p>\n<\/td>\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Mostly solitary, some pairs, small to large groups<\/p>\n<\/td>\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Only two are solitary, all others live in pairs, small to very large groups<\/p>\n<\/td>\n<\/tr>\n<tr class=\"a1-R\">\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Sexual dimorphism<\/strong><\/p>\n<\/td>\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Minimal to none<\/p>\n<\/td>\n<td class=\"a1-C\" style=\"background-color: transparent;vertical-align: middle;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Few taxa have little\/none, many taxa show moderate to high dimorphism<\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<p class=\"import-Normal\">Suborder Haplorrhini is divided into three infraorders: Tarsiiformes, which includes the tarsiers of Asia; Platyrrhini, which includes the monkeys of Central and South America; and Catarrhini, a group that includes the monkeys of Asia and Africa, apes, and humans. According to molecular estimates, tarsiers split from the other haplorrhines close to 70 million years ago, and platyrrhines split from catarrhines close to 46 million years ago (Pozzi Et al. 2014).<\/p>\n<h4 class=\"import-Normal\"><em>Infraorder Tarsiiformes of Asia<\/em><\/h4>\n<figure style=\"width: 188px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image27-1.jpg\" alt=\"Tarsier gripping a branch.\" width=\"188\" height=\"160\" \/><figcaption class=\"wp-caption-text\">Figure 6.23: Tarsiers are the only living representatives of this Infraorder. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Tarsier_Sanctuary,_Corella,_Bohol_(2052878890).jpg\">Tarsier Sanctuary, Corella, Bohol (2052878890)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/46274125@N00\">yeowatzup<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\">CC BY 2.0 License<\/a>.<\/figcaption><\/figure>\n<figure style=\"width: 362px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image28-1.png\" alt=\"Map of Southeast Asia shows distribution of tarsiers.\" width=\"362\" height=\"279\" \/><figcaption class=\"wp-caption-text\">Figure 6.24: Tarsiiformes are found in the tropical forests of multiple islands in Southeast Asia including Sumatra, Borneo, Celebes, and the Philippines. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-4\/\">Infraorder Tarsiiformes of Asia map (Figure 5.22)<\/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>Today, the Infraorder Tarsiiformes includes only one genus, <em>Tarsius <\/em>(Figure 6.23). Tarsiers are small-bodied primates that live in Southeast Asian forests (Figure 6.24) and possess an unusual collection of traits that have led to some debate about their position in the primate taxonomy. They are widely considered members of the haplorrhine group because they share several derived traits with monkeys, apes, and humans, including dry noses, a fovea, not having a tapetum lucidum, and eyes that are more convergent. Tarsiers also have some traits that are more like strepsirrhines and some that are unique. Tarsiers are the only haplorrhine that are specialized vertical clinger leapers, a form of locomotion only otherwise seen in some strepsirrhines. Tarsiers actually get their name because their ankle (tarsal) bones are elongated to provide a lever for vertical clinging and leaping. Tarsiiformes are also small, with most species weighing between 100 and 150 grams. Like strepsirrhines, tarsiers are nocturnal, but because they lack a tapetum lucidum, tarsiers compensate by having enormous eyes. In fact, each eye of a tarsier is larger than its brain. These large eyes allow enough light in for tarsiers to still be able to see well at night without the reflecting layer in their eyes. To protect their large eyes, tarsiers have a partially closed postorbital plate that appears somewhat intermediate between the postorbital bar of strepsirrhines and the full postorbital closure of other haplorrhines (Figure 6.25). Tarsiers have different dental formulas on their upper and lower teeth. On the top, the dental formula is 2:1:3:3, but on the bottom it is 1:1:3:3. Other unusual traits of tarsiers include having two grooming claws on each foot and the ability to rotate their heads around 180 degrees, a trait useful in locating insect prey. The tarsier diet is considered <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1416\">faunivorous<\/a> <\/strong>because it consists entirely of animal matter, making them the only primate not to eat any vegetation. They are only one of two living haplorrhines to be solitary, the other being the orangutan. Most tarsiers are not sexually dimorphic, like strepsirrhines, although males of a few species are slightly larger than females.<\/p>\n<figure style=\"width: 486px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image29-1.jpg\" alt=\"Front view of tarsier skull.\" width=\"486\" height=\"323\" \/><figcaption class=\"wp-caption-text\">Figure 6.25: Skull of a tarsier showing very large eye sockets and partially closed postorbital plates. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Tarsier_skull.jpg\">Tarsier skull<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/65438265@N00\">Andrew Bardwell<\/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\">Two alternative classifications have emerged due to the unusual mix of traits that tarsiers have. Historically, tarsiers were grouped with lemurs, lorises, and galagos into a suborder called Prosimii. This classification was based on tarsiers, lemurs, lorises, and galagos all having grooming claws and similar lifestyles. Monkeys, apes, and humans were then separated into a suborder called the Anthropoidea. These suborder groupings were based on <em>grade<\/em> rather than <em>clade<\/em>. Today, most people use Suborders Strepsirrhini and Haplorrhini, which are clade groupings based on the derived traits that tarsiers share with monkeys, apes, and humans. The Strepsirrhini\/Haplorrhini dichotomy is also supported by the genetic evidence that indicates tarsiers are more closely related to monkeys, apes, and humans (Jameson Et al. 2011). Figure 6.26 summarizes the unusual mix of traits seen in tarsiers.<\/p>\n<table class=\"aligncenter\" style=\"width: 468pt\">\n<caption>Figure 6.26: Tarsiers at a glance: Tarsiers have a mix of traits that lead to debate about their classification. While they have some unique characteristics, they also have traits that superficially resemble strepsirrhines, and many derived traits shared with haplorrhines. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-4\/\">Tarsiers at a glance table (Figure 5.24)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Stephanie Etting 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>\n<td class=\"a2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Like Strepsirrhini<\/strong><\/p>\n<\/td>\n<td class=\"a2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Unique<\/strong><\/p>\n<\/td>\n<td class=\"a2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Like Haplorrhini<\/strong><\/p>\n<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"a2-R\">\n<td class=\"a2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Very small<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Nocturnal<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Highly insectivorous<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Solitary<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Vertical clinger-leapers<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Little\/no sexual dimorphism<\/p>\n<\/td>\n<td class=\"a2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Two grooming claws<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">2:1:3:3\/1:1:3:3 dental formula<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Do not eat vegetation<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Can rotate their heads nearly 180 degrees<\/p>\n<\/td>\n<td class=\"a2-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Almost full PO closure<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">More convergent eyes<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">No tapetum lucidum<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">No rhinarium<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Genetic evidence<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Fovea<\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<h4 class=\"import-Normal\"><em>Infraorder Platyrrhini of Central and South America<\/em><\/h4>\n<figure id=\"attachment_179\" aria-describedby=\"caption-attachment-179\" style=\"width: 329px\" class=\"wp-caption alignleft\"><img class=\"wp-image-165\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image30-e1686352290168.png\" alt=\"Map of South America shows where platyrrhines live.\" width=\"329\" height=\"324\" \/><figcaption id=\"caption-attachment-179\" class=\"wp-caption-text\">Figure 6.27: Geographic distribution of the platyrrhines across the southern part of Central America and the tropical and termporate regions of South America. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-4\/\">Infraorder Platyrrhini map<\/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>The platyrrhines are the only nonhuman primates in Central and South America (Figure 6.27) and so, like the lemurs of Madagascar, have diversified into a variety of forms in the absence of competition. Infraorder Platyrrhini get their name from their distinctive nose shape. \u201cPlaty\u201d means flat and \u201crhini\u201d refers to noses, and, indeed, platyrrhines have noses that are flat and wide, with nostrils that are far apart, facing outward, and usually round in shape (Figure 6.28). This nose shape is very different from what we see in catarrhines.<\/p>\n<p>On average, platyrrhines are smaller and less sexually dimorphic than catarrhines, and they have retained the more ancestral primate dental formula of 2:1:3:3. Platyrrhines are all highly arboreal, whereas many catarrhines spend significant time on the ground. The monkeys in Central and South America also differ in having less well-developed vision. This is reflected in the wiring in the visual system of the brain as well as in their <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1418\">polymorphic color vision<\/a><\/strong>. The genes that enable individuals to distinguish reds and yellows from blues and greens are on the X chromosome. Different genes code for being able to see different wavelengths of light so to distinguish between them you need to be heterozygous for seeing colour. The X chromosomes of platyrrhines each carry the genes for seeing one wavelength, so male platyrrhines (with only one X chromosome) are always dichromatic. Female platyrrhines can be dichromatic (if they are homozygous for one version of the colour vision gene) or trichromatic (if they are heterozygous) (Kawamura Et al. 2012). We currently know of two exceptions to this pattern among platyrrhines. Nocturnal owl monkeys are <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1420\">monochromatic<\/a><\/strong>, meaning that they cannot distinguish any colours. The other exception are howler monkeys, which have evolved to have two colour vision genes on each X chromosome. This means that both male and female howler monkeys are able to see reds and yellows. By contrast, catarrhine males and females are all trichromatic.<\/p>\n<figure style=\"width: 279px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image31-1.jpg\" alt=\"White-faced capuchin monkey.\" width=\"279\" height=\"190\" \/><figcaption class=\"wp-caption-text\">Figure 6.28: A capuchin monkey demonstrating a typical platyrrhine nose shape with round nostrils pointing outward on a flat nose. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:CARABLANCA_-_panoramio.jpg\">CARABLANCA - panoramio<\/a> by Manuel Velazquez is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/3.0\/legalcode\">CC BY 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">Platyrrhines include the smallest of the monkeys, the marmosets and tamarins (Figure 6.29), all of which weigh less than one kilogram and live in cooperative family groups, wherein usually only one female reproduces and everyone else helps carry and raise the offspring. They are unusual primates in that they regularly produce twins. Marmosets and tamarins largely eat gums and saps, so these monkeys have evolved claw-like nails that enable them to cling to the sides of tree trunks like squirrels as well as special teeth that allow them to gnaw through bark. Except for the Goeldi\u2019s monkey, these small monkeys have one fewer molar than other platyrrhines, giving them a dental formula of 2:1:3:2.<\/p>\n<figure style=\"width: 428px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image32-1.jpg\" alt=\"Six marmoset and tamarin species.\" width=\"428\" height=\"470\" \/><figcaption class=\"wp-caption-text\">Figure 6.29: Clockwise from top right: golden-headed lion tamarin, pygmy marmoset, Goeldi\u2019s monkey, bare-eared marmoset, emperor tamarin, and common marmoset. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Callitrichinae_genus.jpg\">Callitrichinae genus<\/a> by Miguelrangeljr is a collective work under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/deed.en\">CC BY-SA 3.0 License<\/a>. [Includes <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Wei%C3%9Fb%C3%BCschelaffe_(Callithrix_jacchus).jpg\">Wei\u00dfb\u00fcschelaffe_(Callithrix_jacchus)<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Raymond\">Raymond<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/\">CC BY-SA 4.0<\/a>;\u00a0 <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Leontopithecus_chrysomelas_(portrait).jpg\">Leontopithecus chrysomelas (portrait)<\/a> by Hans Hillewaert, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/\">CC BY-SA 4.0<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Emperor_Tamarin_portrait_2_edit1.jpg\">Emperor_Tamarin_portrait_2_edit1<\/a> by <a href=\"https:\/\/sites.google.com\/site\/thebrockeninglory\/\">Brocken Inaglory<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/\">CC BY-SA 4.0<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Dv%C3%A6rgsilkeabe_Callithrix_pygmaea.jpg\">Dv\u00e6rgsilkeabe_Callithrix_pygmaea<\/a> by Malene Thyssen (User <a href=\"https:\/\/da.wikipedia.org\/wiki\/User:Malene\">Malene<\/a>), <a href=\"https:\/\/en.wikipedia.org\/wiki\/GNU_Free_Documentation_License\">GNU Free Documentation License<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Mico_argentatus_(portrait).jpg\">Mico_argentatus_(portrait)<\/a> by Hans Hillewaert, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/\">CC BY-SA 4.0<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Titi_Monkey.jpg\">Titi Monkey<\/a> by Jeff Kubina, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/deed.en\">CC BY-SA 2.0<\/a>].]<\/figcaption><\/figure>\n<p class=\"import-Normal\">The largest platyrrhines are a family that include spider monkeys, woolly spider monkeys, woolly monkeys, and howler monkeys (Figure 6.30). These monkeys can weigh up to 9\u201315 kg and have evolved prehensile tails that can hold their entire body weight. It is among this group that we see semi-brachiators, like the spider monkey (see Figure 6.13). To make them more efficient in this form of locomotion, spider monkeys evolved to not have thumbs so that their hands work more like hooks that can easily let go of branches while swinging. Howler monkeys are another well-known member of this group, earning their name due to their loud calls, which can be heard miles away. To make these loud vocalizations, howler monkeys have a specialized vocal system that includes a large larynx and hyoid bone. Howler monkeys are the most folivorous of the platyrrhines and are known for spending a large portion of their day digesting their food.<\/p>\n<figure style=\"width: 458px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image33-1.jpg\" alt=\"Four platyrrhine species.\" width=\"458\" height=\"457\" \/><figcaption class=\"wp-caption-text\">Figure 6.30: Clockwise from top right: howler monkey, woolly monkey, woolly spider monkey, and spider monkey. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Atelidae_Family.jpg\">Atelidae Family<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Miguelrangeljr\">Miguelrangeljr<\/a> is a collective work under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>. [Includes <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Ateles_marginatus_(Sao_Paulo_zoo).jpg\">Ateles marginatus (Sao Paulo zoo)<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Miguelrangeljr\">Miguelrangeljr<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Alouatta_caraya_male.JPG\">Alouatta caraya male<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Miguelrangeljr\">Miguelrangeljr<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Lagothrix_lagotricha_(walking).jpg\">Lagothrix lagotricha (walking)<\/a> by Hans Hillewaert, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Brachyteles_hypoxanthus2.jpg\">Brachyteles hypoxanthus2<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/42956474@N04\/with\/4133258867\/\">Paulo B. Chaves<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\">CC BY 2.0<\/a>.]<\/figcaption><\/figure>\n<p class=\"import-Normal\">There are many other monkeys in Central and South America, including the gregarious capuchins (see Figure 6.28) and squirrel monkeys, the pair-living titi monkeys, and the nocturnal owl monkeys. There are also the seed-eating saki monkeys and uakaris. In many areas across Central and South America, multiple species of platyrrhines share the forests, with some even traveling together in association. According to molecular evidence, the diversity of platyrrhines that we see today seems to have originated about 25 million years ago (Schneider &amp; Sampaio 2015). Figure 6.31 summarizes the key traits of platyrrhines relative to the other infraorders of Haplorrhini.<\/p>\n<table class=\"aligncenter\" style=\"width: 468pt\">\n<caption>Figure 6.31: Platyrrhini at a glance: Summary of the key traits we use to distinguish platyrrhines. Traits indicated with an * are those with exceptions detailed in the text. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-4\/\">Platyrrhini at a glance table (Figure 5.29)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Stephanie Etting is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>.<\/caption>\n<tbody>\n<tr class=\"a3-R\">\n<td class=\"a3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Platyrrhini traits<\/strong><\/p>\n<\/td>\n<\/tr>\n<tr class=\"a3-R\">\n<td class=\"a3-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Flat nose with rounded nostrils pointing to the side<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Highly arboreal<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Less sexually dimorphic on average<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">2:1:3:3 dental formula*<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Polymorphic colour vision*<\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<h4 class=\"import-Normal\"><em>Infraorder Catarrhini of Asia and Africa <\/em><\/h4>\n<figure style=\"width: 191px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image34.jpg\" alt=\"Wolf\u2019s guenon.\" width=\"191\" height=\"287\" \/><figcaption class=\"wp-caption-text\">Figure 6.32: A Wolf\u2019s guenon demonstrating a typical catarrhine nose with teardrop-shaped nostrils close together and pointed downward. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Wolf's_Guenon_Picking_Up_Food_(19095137693).jpg\">Wolf's Guenon Picking Up Food (19095137693)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/8749778@N06\">Eric Kilby<\/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\">Infraorder Catarrhini includes Superfamily Cercopithecoidea (the monkeys of Africa and Asia) and Superfamily Hominoidea (apes and humans). Nonhuman catarrhines are found all over Africa and South and Southeast Asia, with some being found as far north as Japan. The most northerly and southerly catarrhines are cercopithecoid monkeys. In contrast, apes are less tolerant of drier, more seasonal environments and so have a relatively restricted geographic range.<\/p>\n<p class=\"import-Normal\">Relative to other haplorrhine infraorders, catarrhines are distinguished by several characteristics. Catarrhines have a distinctive nose shape, with teardrop-shaped nostrils that are close together and point downward (Figure 6.32) and one fewer premolar than most other primates, giving us a dental formula of 2:1:2:3 (Figure 6.33). On average, catarrhines are the largest and most sexually dimorphic of all primates. Gorillas are the largest living primates, with males weighing up to 220 kg. The most sexually dimorphic of all primates are mandrills. Mandrill males not only have much more vibrant colouration than mandrill females but also have larger canines and can weigh up to three times more (Setchell Et al. 2001). The larger body size of catarrhines is related to the more terrestrial lifestyle of many members of this infraorder. In fact, the most terrestrial of living primates can be found in this group. Among all primates, vision is the most developed in catarrhines. Catarrhines independently evolved the same adaptation as howler monkeys in having each X chromosome with genes to distinguish both reds and yellows, so all male and female catarrhines are trichromatic, which is useful for these diurnal primates.<\/p>\n<figure style=\"width: 632px\" class=\"wp-caption aligncenter\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image35-1.jpg\" alt=\"Platyrrhine, cercopithecoid, and hominoid mandibles.\" width=\"632\" height=\"331\" \/><figcaption class=\"wp-caption-text\">Figure 6.33: Catarrhines have two premolars whereas most other primate taxa (including platyrrhini) have three premolars. This image also shows one of the derived traits of cercopithecoids, their bilophodont molars, which differ from the more ancestral Y-5 molars of apes and humans. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-4\/\">Platyrrhini vs. Catarrhini dentition<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Stephanie Etting 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:\/\/animaldiversity.org\/accounts\/Animalia\/specimens\/collections\/contributors\/phil_myers\/ADW_mammals\/specimens\/Primates\/Cebus_apella\/lower_dorsal2216\/?start=135;f=subject::specimen::lower%20jaw\">Cebus apella (brown capuchin)<\/a> at Animal Diversity Web by <a href=\"https:\/\/animaldiversity.org\/collections\/contributors\/phil_myers\/ADW_mammals\/specimens\/Primates\/Hylobates_syndactylus\/lower_dorsal0097\/\">Phil Myers<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/\">CC BY-NC-SA 3.0<\/a>; <a href=\"https:\/\/animaldiversity.org\/accounts\/Animalia\/specimens\/collections\/contributors\/phil_myers\/ADW_mammals\/specimens\/Primates\/Lophocebus_albigena\/lower_dorsal2060\/?start=525;f=subject::specimen::lower%20jaw\">Lophocebus albigena (gray-cheeked mangaby)<\/a> at Animal Diversity Web by <a href=\"https:\/\/animaldiversity.org\/collections\/contributors\/phil_myers\/ADW_mammals\/specimens\/Primates\/Hylobates_syndactylus\/lower_dorsal0097\/\">Phil Myers<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/3.0\/\">CC BY-NC-SA 3.0<\/a>; <a href=\"https:\/\/animaldiversity.org\/accounts\/Primates\/specimens\/collections\/contributors\/phil_myers\/ADW_mammals\/specimens\/Primates\/Hylobates_syndactylus\/lower_dorsal0097\/?start=105;f=subject::specimen::lower%20jaw\">Symphalangus syndactylus (siamang)<\/a> at Animal Diversity Web by <a href=\"https:\/\/animaldiversity.org\/collections\/contributors\/phil_myers\/ADW_mammals\/specimens\/Primates\/Hylobates_syndactylus\/lower_dorsal0097\/\">Phil Myers<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/3.0\/\">CC BY-NC-SA 3.0<\/a>.]<\/figcaption><\/figure>\n<p class=\"import-Normal\">The two superfamilies of catarrhines\u2014Superfamily Cercopithecoidea, the monkeys of Africa and Asia, and Superfamily Hominoidea, which includes apes and humans\u2014are believed to have split about 32 million years ago based on molecular evidence (Pozzi Et al. 2014). This fits with the fossil record, which shows evidence of these lineages by about 25 million years ago (see Chapters 8-9).<\/p>\n<h4 class=\"import-Normal\"><em>Superfamily Cercopithecoidea of Africa and Asia<\/em><\/h4>\n<figure style=\"width: 285px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image36-1.jpg\" alt=\"Pinkish ischial callosities on a crested black macaque.\" width=\"285\" height=\"214\" \/><figcaption class=\"wp-caption-text\">Figure 6.34: The second derived trait of cercopithecoids are their ischial callosities, shown here on a crested black macaque. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Sulawesi_trsr_DSCN0572_v1.JPG\">Sulawesi trsr DSCN0572 v1<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Shankar_Raman\">T. R. Shankar Ramanis<\/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>Compared to hominoids, cercopithecoids have an ancestral quadrupedal body plan with two key derived traits. The first derived trait of cercopithecoids is their <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1422\">bilophodont<\/a> <\/strong>molars (\u201cbi\u201d meaning two, \u201cloph\u201d referring to ridge, and \u201cdont\u201d meaning tooth). If you refer back to Figure 6.33, you will see how the molars of cercopithecoids have four cusps arranged in a square pattern and have two ridges connecting them. It is thought that this molar enabled these monkeys to eat a wide range of foods, thus allowing them to live in habitats that apes cannot. The other key derived trait that all cercopithecoids share is having <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1424\">ischial callosities<\/a> <\/strong>(Figure 6.34). The ischium is the part of your pelvis that you are sitting on right now (see Appendix A: Osteology). In cercopithecoids, this part of the pelvis has a flattened surface that, in living animals, has callused skin over it. These function as seat pads for cercopithecoids, who often sit above branches when feeding and resting.<\/p>\n<figure style=\"width: 359px\" class=\"wp-caption alignright\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image37-1.png\" alt=\"Areas of Europe, Asia, Africa, and Australia where cercopithecoids live.\" width=\"359\" height=\"277\" \/><figcaption class=\"wp-caption-text\">Figure 6.35: Geographic distribution of the cercopithecoid monkeys. Catarrhines have the widest geographic distribution due to the success of cercopithecoids who are found all across subsaharan Africa and southern Asia. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-4\/\">Superfamily Cercopithecoidea map (Figure 5.33)<\/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>Cercopithecoid monkeys are the most geographically widespread group of nonhuman primates (Figure 6.35). Since their divergence from hominoids, this monkey group has increased in numbers and diversity due, in part, to their fast reproductive rates. On average, cercopithecoids will reproduce every one to two years, whereas hominoids will reproduce once every four to nine years, depending on the taxon.<\/p>\n<figure style=\"width: 180px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image38-1.jpg\" alt=\"Two silver leaf monkeys hold orange-haired infants.\" width=\"180\" height=\"240\" \/><figcaption class=\"wp-caption-text\">Figure 6.36: Silver leaf monkey infants are born with orange fur, dramatically contrasting the adult coat colour of their mothers. After a few months, the infants gradually change colour to that of their parents. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Silverleaf_Monkey_(Kuala_Lumpur).jpg\">Silverleaf Monkey (Kuala Lumpur)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/10815162@N07\">Andrea Lai<\/a> from Auckland, New Zealand, is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/legalcode\">CC BY 2.0 License.<\/a><\/figcaption><\/figure>\n<p>Cercopithecoidea is split into two groups, the leaf monkeys and the cheek-pouch monkeys. Both groups coexist in Asia and Africa; however, the majority of leaf monkey species live in Asia with only a few taxa in Africa. In contrast, only one genus of cheek-pouch monkey lives in Asia, and all the rest of them in Africa. As you can probably guess based on their names, the two groups differ in terms of diet. Leaf monkeys are primarily folivores, with some species eating a significant amount of seeds. Cheek-pouch monkeys tend to be more frugivorous or omnivorous, with one taxon, geladas, eating primarily grasses. The two groups also differ in some other interesting ways. Leaf monkeys tend to produce infants with <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1426\">natal coats<\/a><\/strong>\u2014infants whose fur is a completely different colour from their parents (Figure 6.36). Leaf monkeys are also known for having odd noses (Figure 6.37), and so they are sometimes called \u201codd-nosed monkeys.\u201d Cheek-pouch monkeys are able to pack food into their cheek pouches (Figure 6.38), thus allowing them to move to a location safe from predators or aggressive individuals of their own species where they can eat in peace.<\/p>\n<figure style=\"width: 408px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image39.jpg\" alt=\"Male proboscis monkey.\" width=\"408\" height=\"272\" \/><figcaption class=\"wp-caption-text\">Figure 6.37: Proboscis monkeys are one of several \u201codd-nosed\u201d leaf monkeys. Male proboscis monkeys, like this one, have large, pendulous noses, while females have much smaller noses. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Proboscis_monkey_(Nasalis_larvatus)_male_head.jpg\">Proboscis monkey (Nasalis larvatus) male head<\/a> by <a href=\"https:\/\/www.sharpphotography.co.uk\/\">Charles J Sharp<\/a> creator QS:P170,Q54800218 is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/legalcode\">CC BY-SA 4.0 License<\/a>.<\/figcaption><\/figure>\n<figure style=\"width: 414px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image40-1.jpg\" alt=\"Bonnet macaque with full cheek pouches.\" width=\"414\" height=\"275\" \/><figcaption class=\"wp-caption-text\">Figure 6.38: This bonnet macaque has filled its cheek pouches with food, an adaptation that is useful in transporting food to a safer location to eat. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Bonnet_macaque_DSC_0893.jpg\">Bonnet macaque DSC 0893<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Shankar_Raman\">T. R. Shankar Raman<\/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\"><em>Superfamily Hominoidea of Africa and Asia<\/em><\/h4>\n<figure style=\"width: 438px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image41.png\" alt=\"Areas of Europe, Asia, Africa, and Australia where hominoidea live.\" width=\"438\" height=\"339\" \/><figcaption class=\"wp-caption-text\">Figure 6.39: Geographic distribution of apes across Central and West Africa and Southeast Asia. Hominoids overlap geographically with cercopithecoid monkeys but have a lower tolerance for seasonal environments and so are found only in tropical forests across these regions. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-4\/\">Superfamily Hominoidea map (Figure 5.38)<\/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\">Superfamily Hominoidea of Africa and Asia (Figure 6.39) includes the largest of the living primates: apes and humans. Whereas cercopithecoid monkeys have bilophodont molars, hominoids have the more ancestral <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1428\">Y-5 molars<\/a><\/strong>, which feature five cusps separated by a \u201cY\u201d-shaped groove pattern (see Figure 6.33). The Y-5 molar was present in the common ancestors of hominoids and cercopithecoids, thus it is the more ancestral molar pattern of the two. Hominoids differ the most from other primates in our body plans, due to the unique form of locomotion that hominoids are adapted for: brachiation (Figure 6.40).<\/p>\n<p>To successfully swing below branches, many changes to the body needed to occur. Hominoid arms are much longer than the legs to increase reach, and the lower back is shorter and less flexible to increase control when swinging. The torso, shoulders, and arms of hominoids have evolved to increase range of motion and flexibility (see again Figure 6.12). The clavicle, or collar bone, is longer to stabilize the shoulder joint out to the side, thus enabling us to rotate our arms 360 degrees. Hominoid rib cages are wider side to side and shallower front to back than those of cercopithecoids and we do not have tails, as tails are useful for balance when running on all fours but generally not useful while swinging. Hominoids also have modified ulnae, one of the two bones in the forearm (see Appendix A: Osteology). At the elbow end of the ulna, hominoids have a short <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1430\">olecranon process<\/a><\/strong>, which allows for improved extension in our arms. At the wrist end of the ulna, hominoids have a short <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1432\">styloid process<\/a><\/strong>, which enables us to have very flexible wrists, a trait critical for swinging. Both the olecranon process and styloid process are long in quadrupedal animals who carry much of their weight on their forelimbs when traveling and who therefore need greater stability rather than flexibility in those joints.<\/p>\n<table class=\"aligncenter\" style=\"width: 468pt\">\n<caption>Figure 6.40: Quadrupedalism vs. brachiation: Summary of the key anatomical differences between a quadrupedal primate and one adapted for brachiation. To view these traits using photos of bones, check out the interactive skeletal websites in \u201cFurther Explorations\u201d below. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-4\/\">Quadrupedalism vs. Brachiation table (Figure 5.39)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Stephanie Etting 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>\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">\n<\/td>\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Quadrupedalism<\/strong><\/p>\n<\/td>\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Brachiation<\/strong><\/p>\n<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"a4-R\">\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Arm length vs. leg length<\/strong><\/p>\n<\/td>\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">About equal<\/p>\n<\/td>\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Arms are longer<\/p>\n<\/td>\n<\/tr>\n<tr class=\"a4-R\">\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Shoulder position<\/strong><\/p>\n<\/td>\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">More on the front<\/p>\n<\/td>\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Out to the side<\/p>\n<\/td>\n<\/tr>\n<tr class=\"a4-R\">\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Ribcage shape<\/strong><\/p>\n<\/td>\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Deep front-to-back<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Narrow side-to-side<\/p>\n<\/td>\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Shallow front-to-back<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Wide side-to-side<\/p>\n<\/td>\n<\/tr>\n<tr class=\"a4-R\">\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Length of lower back<\/strong><\/p>\n<\/td>\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Long<\/p>\n<\/td>\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Short<\/p>\n<\/td>\n<\/tr>\n<tr class=\"a4-R\">\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Collar bone length<\/strong><\/p>\n<\/td>\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Short<\/p>\n<\/td>\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Long<\/p>\n<\/td>\n<\/tr>\n<tr class=\"a4-R\">\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Ulnar olecranon process<\/strong><\/p>\n<\/td>\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Long<\/p>\n<\/td>\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Short<\/p>\n<\/td>\n<\/tr>\n<tr class=\"a4-R\">\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Ulnar styloid process<\/strong><\/p>\n<\/td>\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Long<\/p>\n<\/td>\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Short<\/p>\n<\/td>\n<\/tr>\n<tr class=\"a4-R\">\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Tail<\/strong><\/p>\n<\/td>\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Short to long<\/p>\n<\/td>\n<td class=\"a4-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">None<\/p>\n<\/td>\n<\/tr>\n<tr>\n<td><\/td>\n<td><\/td>\n<td><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<p class=\"import-Normal\">Apes and humans also differ from other primates in behaviour and life history characteristics. Hominoids all seem to show some degree of female dispersal at sexual maturity but, as you will learn in Chapter 7, it is more common that males leave. Some apes show males dispersing in addition to females, but the hominoid tendency for female dispersal is a bit unusual among primates. Our superfamily is also characterized by the most extended life histories of all primates. All members of this group take a long time to grow and reproduce much less frequently compared to cercopithecoids. The slow pace of this life history is likely related to why hominoids have decreased in diversity since they first evolved. Figure 6.41 summarizes the key traits of Infraorder Catarrhini and its two superfamilies. Today, there are only five types of hominoids left: gibbons and siamangs, orangutans, gorillas, chimpanzees and bonobos, and humans.<\/p>\n<table class=\"aligncenter\">\n<caption>Figure 6.41a: Catarrhini at a glance: Summary of key traits of the Infraorder Catarrhini. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-4\/\">Catarrhini at a glance (Figure 5.40)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Stephanie Etting is a collective work under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. [Includes <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Duskyleafmonkey1.jpg\">Duskyleafmonkey1<\/a> by <a href=\"https:\/\/www.the-ninth.com\/about\">Robertpollai<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by\/3.0\/at\/deed.en\">CC BY 3.0 AT<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Male_Bornean_Orangutan_-_Big_Cheeks.jpg\">Male Bornean Orangutan - Big Cheeks<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/8749778@N06\">Eric Kilby<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/legalcode\">CC BY-SA 2.0<\/a>.]<\/caption>\n<thead>\n<tr style=\"height: 22pt\">\n<td class=\"a5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\" colspan=\"2\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong>Infraorder Catarrhini<\/strong><\/p>\n<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"a5-R\" style=\"height: 22pt\">\n<td class=\"a5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\" colspan=\"2\">\n<p class=\"import-Normal\" style=\"text-align: center\">Downward facing, tear-drop shaped nostrils, close together<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Arboreal and more terrestrial taxa<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">On average, largest primates<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">On average, most sexually dimorphic taxonomic group<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">2:1:2:3 dental formula<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">All trichromatic<\/p>\n<\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<table class=\"aligncenter\" style=\"width: 468pt\">\n<caption>Figure 6.41b: Characteristics used to distinguish between the two Catarrhini superfamilies. Credit: <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/chapter\/__unknown__-4\/\">Catarrhini at a glance (Figure 5.40)<\/a> original to <a href=\"https:\/\/pressbooks-dev.oer.hawaii.edu\/explorationsbioanth\/\">Explorations: An Open Invitation to Biological Anthropology<\/a> by Stephanie Etting is a collective work under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/\">CC BY-NC 4.0 License<\/a>. [Includes <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Duskyleafmonkey1.jpg\">Duskyleafmonkey1<\/a> by <a href=\"https:\/\/www.the-ninth.com\/about\">Robertpollai<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by\/3.0\/at\/deed.en\">CC BY 3.0 AT<\/a>; <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Male_Bornean_Orangutan_-_Big_Cheeks.jpg\">Male Bornean Orangutan - Big Cheeks<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/8749778@N06\">Eric Kilby<\/a>, <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/legalcode\">CC BY-SA 2.0<\/a>.]<\/caption>\n<thead>\n<tr>\n<td class=\"a5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong><img class=\"alignnone\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image42-1.jpg\" alt=\"Dusky leaf monkey\" width=\"248\" height=\"186\" \/><\/strong><strong>Superfamily Cercopithecoidea<\/strong><\/p>\n<\/td>\n<td class=\"a5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\"><strong><img class=\"alignnone\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image43-1.jpg\" alt=\"Orangutan\" width=\"274\" height=\"183\" \/><\/strong><strong>Superfamily Hominoidea<\/strong><\/p>\n<\/td>\n<\/tr>\n<\/thead>\n<tbody>\n<tr class=\"a5-R\">\n<td class=\"a5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Wide geographic distribution<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Bilophodont molars<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Ischial callosities<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Reproduce every 1\u20132 years<\/p>\n<\/td>\n<td class=\"a5-C\" style=\"background-color: transparent;padding: 5pt 5pt 5pt 5pt;border: solid #000000 1pt\">\n<p class=\"import-Normal\" style=\"text-align: center\">Tropical forests of Africa and Asia<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Y-5 molars<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Adaptations for brachiation<\/p>\n<p class=\"import-Normal\" style=\"text-align: center\">Reproduce every 4\u20139 years<\/p>\n<\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<h4 class=\"import-Normal\"><em>Family <\/em>Hylobatidae <em>of Southeast Asia<\/em><\/h4>\n<figure style=\"width: 441px\" class=\"wp-caption alignleft\"><img class=\"\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image44-1.jpg\" alt=\"Siamang with outstretched arms.\" width=\"441\" height=\"294\" \/><figcaption class=\"wp-caption-text\">Figure 6.42: Siamangs are the largest of the Hylobatidae family. They are all black with a throat sac that can become inflated to give out loud calls. Credit: <a href=\"https:\/\/www.flickr.com\/photos\/suneko\/373310729\/\">Shout (373310729)<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/suneko\/\">su neko<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\">CC BY-SA 2.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">The number of genera in this group has been changing in recent years, but the taxa broadly encompasses gibbons and siamangs. Both are found across Southeast Asian tropical forests. Gibbons weigh, on average, about 13 pounds and tend to be more frugivorous, whereas siamangs are larger than gibbons and also more folivorous. Unlike the larger-bodied apes (orangutans, chimps, bonobos, and gorillas) who make nests to sleep in every night, gibbons and siamangs will develop callused patches on their ischium resembling ischial callosities. Gibbon species are quite variable in their colouration and markings, while siamangs are all black with big throat sacs that are used in their exuberant vocalizations (Figure 6.42). Both gibbons and siamangs live in pairs with very little sexual dimorphism, although males and females do differ in colouration in some gibbon species.<\/p>\n<h4 class=\"import-Normal\">Pongo<em> of Southeast Asia<\/em><\/h4>\n<p class=\"import-Normal\">The Genus <em>Pongo <\/em>refers to orangutans. These large red apes are found in Southeast Asia, with the two well-known species each living on the islands of Borneo and Sumatra. A third, very rare species, was recently discovered in Southern Sumatra (Nater Et al. 2017). Orangutans are highly frugivorous but will supplement their diet with leaves and bark when fruit is less available. As mentioned earlier, orangutans are the only diurnal, solitary taxon among primates and are extremely slow to reproduce, producing only one offspring about every seven to nine years. They are highly sexually dimorphic (Figure 6.43 a &amp; b), with fully developed, \u201cflanged\u201d males being approximately twice the size of females. These males have large throat sacs; long, shaggy coats; and cheek flanges. The skulls of male orangutans often feature a <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1434\">sagittal crest<\/a><\/strong>, which is believed to function as additional attachment area for chewing muscles as well as a trait used in sexual competition (Balolia, Soligo, &amp; Wood 2017). An unusual feature of orangutan biology is <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1436\">male bimaturism<\/a><\/strong>. Male orangutans are known to delay maturation until one of the more dominant, flanged males disappears. The males that delay maturation are called \u201cunflanged\u201d males, and they can remain in this state for their entire life. Unflanged males resemble females in their size and appearance and will sneak copulations with females while avoiding the bigger, flanged males. Flanged and unflanged male orangutans represent alternative reproductive strategies, both of which successfully produce offspring (Utami Et al. 2002).<\/p>\n<p>&nbsp;<\/p>\n<\/div>\n<figure id=\"attachment_181-4\" aria-describedby=\"caption-attachment-181-4\" style=\"width: 1900px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-180 size-full\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/5.43.jpg\" alt=\"a. Female orangutan with infant. b. Male orangutan in a tree.\" width=\"1900\" height=\"800\" \/><figcaption id=\"caption-attachment-181-4\" class=\"wp-caption-text\">Figure 6.43: (a) A female orangutan eating fruit with her infant nearby and (b) a flanged adult male eating leaves. Male orangutans are about twice the size of females and have a longer coat length, cheek flanges, and throat sac. Credit: 6.43a. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Orang_Utan_(Pongo_pygmaeus)_female_with_baby_(8066259067).jpg\">Orang Utan (Pongo pygmaeus) female with baby (8066259067)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/65695019@N07\">Bernard DUPONT<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/2.0\/\">CC BY-SA 2.0 Licence<\/a>. 6.43b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Orangutan_-Zoologischer_Garten_Berlin-8a.jpg\">Orangutan -Zoologischer Garten Berlin-8a<\/a> by <a href=\"https:\/\/www.flickr.com\/photos\/37088680@N03\">David Forsman<\/a> is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/2.0\/\">CC BY 2.0 License<\/a>.<\/figcaption><\/figure>\n<div class=\"learning-objectives\">\n<h4 class=\"import-Normal\">Gorilla <em>of Africa<\/em><\/h4>\n<p class=\"import-Normal\">There are several species of gorillas that can be found across Central Africa. Gorilla males, like orangutan males, are about twice the size of female gorillas (Figures 6.44a &amp; b). When on the ground, gorillas use a form of quadrupedalism called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1438\">knuckle-walking<\/a><\/strong>, wherein the fingers are curled under and the weight is carried on the knuckles. Male gorillas have a large sagittal crest and large canines compared with females. Adult male gorillas are often called \u201csilverbacks\u201d because when they reach about twelve to thirteen years old, the hair on their backs turns silvery grey. Gorillas typically live in groups of one male and several females. Gorillas are considered folivorous, although some species can be more frugivorous depending on fruit seasonality (Remis 1997).<\/p>\n<\/div>\n<figure id=\"attachment_181-5\" aria-describedby=\"caption-attachment-181-5\" style=\"width: 1900px\" class=\"wp-caption aligncenter\"><img class=\"wp-image-181 size-full\" src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/5.44.jpg\" alt=\"a. Female gorilla with offspring. b. Male gorilla.\" width=\"1900\" height=\"800\" \/><figcaption id=\"caption-attachment-181-5\" class=\"wp-caption-text\">Figure 6.44: (a) A female gorilla with her two offspring and (b) a silverback adult male. Male gorillas are about twice the size of females. They also differ from females in having a large sagittal crest and a silver back, which appears as they mature. Credit: 6.44a. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Enzo_naomi_echo.jpg\">Enzo naomi echo<\/a> by Zoostar is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by\/3.0\/legalcode\">CC BY 3.0 License<\/a>. 6.44b. <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Male_gorilla_in_SF_zoo.jpg\">Male gorilla in SF zoo<\/a> by Brocken Inaglory is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/legalcode\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<div class=\"learning-objectives\">\n<h4 class=\"import-Normal\">Pan<em> of Africa<\/em><\/h4>\n<figure style=\"width: 252px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image49-1.jpg\" alt=\"Bonobo looks away from the camera.\" width=\"252\" height=\"222\" \/><figcaption class=\"wp-caption-text\">Figure 6.45: Bonobo (Pan paniscus). You can see the distinctive hair-part on this bonobo. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Bonobo_male_Jasongo_15yo_Twycross_582a_(2014_11_14_01_04_18_UTC).jpg\">Bonobo male Jasongo 15yo Twycross 582a (2014 11 14 01 04 18 UTC)<\/a> by William H. Calvin 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>The Genus <em>Pan <\/em>includes two species: <em>Pan troglodytes <\/em>(the common chimpanzee) and <em>Pan paniscus <\/em>(the bonobo). These species are separated by the Congo River, with chimpanzees ranging across West and Central Africa and bonobos located in a restricted area south of the Congo River. Chimpanzees and bonobos both have broad, largely frugivorous diets.The two species differ morphologically in that bonobos are slightly smaller, have their hair parted down the middle of their foreheads, and are born with dark faces (Figure 6.45). In contrast, chimpanzees do not have the distinctive parted hair and are born with light faces that darken as they mature (Figure 6.46). Chimpanzees and bonobos live in a grouping called a fission-fusion community, which you will learn more about in Chapter 6. Both species are moderately sexually dimorphic, with males about 20% larger than females. When on the ground, chimpanzees and bonobos knuckle-walk like gorillas do.<\/p>\n<figure style=\"width: 418px\" class=\"wp-caption aligncenter\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image50-1.jpg\" alt=\"Female chimpanzee with offspring in a tree.\" width=\"418\" height=\"278\" \/><figcaption class=\"wp-caption-text\">Figure 6.46: A common chimpanzee (Pan troglodytes) female (centre) and her offspring. Note the pink face of the youngest individual. Bonobos are born with dark-skinned faces, but chimpanzees are born with pink faces that darken with age. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Chimpanzees_in_Uganda_(5984913059).jpg\">Chimpanzees in Uganda (5984913059)<\/a> by <a href=\"https:\/\/www.flickr.com\/people\/57424551@N06\">USAID Africa Bureau<\/a> uploaded by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Elitre\">Elitre<\/a> is in the <a href=\"https:\/\/en.wikipedia.org\/wiki\/Public_domain\">public domain<\/a>.<\/figcaption><\/figure>\n<h4 class=\"import-Normal\">Homo<\/h4>\n<p class=\"import-Normal\">The last member of the Hominoidea to discuss is our own taxon, Genus <em>Homo<\/em>. Later chapters will discuss the many extinct species of <em>Homo<\/em>, but today there is only one living species of <em>Homo, <\/em>our own species, <em>sapiens<\/em>. While it is interesting to focus on how humans differ from apes in many aspects of our morphology, behaviour, and life history, one objective of this chapter, and of biological anthropology in general, is to understand our place in nature. This means looking for aspects of human biology that link us to the taxonomic diversity we have discussed. To that end, here we will focus on similarities humans share with other hominoids.<\/p>\n<p class=\"import-Normal\">Like other hominoids, humans lack a tail and possess upper-body adaptations for brachiation. While our lower body has been modified for a bipedal gait, we are still able to swing from branches and throw a baseball, all thanks to our mobile shoulder joint. Humans, like other hominoids, also have a Y-5 cusp pattern on our molars. All hominoids, including humans, have an extended life history, taking time to grow and develop, and reproducing slowly over a long life span. Lastly, while humans show a great deal of variation across cultures, many human societies show tendencies for female dispersal (Burton Et al. 1996).<\/p>\n<p class=\"import-Normal\">Among the hominoids, humans show particular affinities with other members of the African Clade, <em>Pan <\/em>and <em>Gorilla<\/em>. Humans share over 96% of our DNA with gorillas (Scally Et al. 2012), and over 98% with <em>Pan <\/em>(Ebersberger Et al. 2002). Even without this strong genetic evidence, the African Clade of hominoids share many morphological similarities, including having wide-set eye sockets and backward-sweeping cheekbones. Today, <em>Pan<\/em> and <em>Gorilla<\/em> knuckle-walk when on the ground, and it has been suggested the last common ancestor of chimpanzees, bonobos, gorillas, and humans did as well (Richmond, Begun, &amp; Strait 2001). Further, humans, chimpanzees, and bonobos all live in fission-fusion social groups characterized by shared behaviours, like male cooperation in hunting and territoriality, as well as tool use.<\/p>\n<div class=\"textbox\">\n<h2 class=\"import-Normal\">Special Topic: Primates in Culture and Religion<\/h2>\n<p class=\"import-Normal\">One of the best parts of teaching anthropology for me is getting to spend time watching primates at zoos. What I also find interesting is watching people watch primates. I have very often heard a parent and child walk up to a chimpanzee enclosure and exclaim \u201cLook at the monkeys!\u201d The parent and child often don\u2019t know that a chimpanzee is not a monkey, nor are they likely to know that chimpanzees share more than 98% of their DNA with us. What strikes me as significant is that, although most people do not know the difference between a monkey, an ape, and a lemur, they nonetheless recognize something in the animals as being similar to themselves. In fact, recognition of similarities between humans and other primates is very ancient, dating back far earlier than Linnaeus. For many of us, we only ever get to see primates in zoos and animal parks, but in many areas of the world, humans have coexisted with these animals for thousands of years. In areas where humans and primates have a long, shared history, nonhuman primates often play key roles in creation myths and cultural symbolism.<\/p>\n<p class=\"import-Normal\">Hamadryas baboons feature significantly in Ancient Egyptian iconography. Ancient Egyptian deities and beliefs transformed over time, as did the role of hamadryas baboons. Early on, baboons were thought to represent dead ancestors, and one monkey deity, called Babi or Baba, was thought to feed off of dead souls. Later, baboons became the totem animal for Thoth, the deity of science, writing, wisdom, and measurement, who also wrote the Book of the Dead. Sunbathing hamadryas baboons led ancient Egyptians to associate them with Ra, the sun god, who was the son of Thoth. During mummification, human organs were removed and put into canopic jars, one of which was topped with the head of the baboon-headed god, Hapi. Hamadryas baboons were also often kept as pets, as depicted in hieroglyphics, and occasionally mummified as well.<\/p>\n<p class=\"import-Normal\">On Madagascar, indris and aye-ayes play roles in the creation myths and omens of local people.There are many myths regarding the origins of indris and their relationship to humans, including one where two brothers living in the forest separated, with one brother leaving the forest and becoming a human while the other stayed in the forest to become the indri. Like humans, indris have long legs, no tail, and upright posture. They are considered sacred and are therefore protected. Unfortunately, the aye-aye is not treated with the same reverence. Because of their unusual appearance (see Figure 6.15), aye-ayes are seen as omens of death.They are usually killed when encountered because it is believed that someone will die if an aye-aye points at them.<\/p>\n<p class=\"import-Normal\">In India, monkeys play a key role in the Hindu religion. Hanuman, who resembles a monkey, is a key figure in the Ramayana. Hanuman is thought to be a guardian deity, and so local monkeys like Hanuman langurs and macaques are protected in India (Figure 6.47). In Thailand, where Hinduism is also practiced, the Hindu reverence for monkeys extends to \u201cmonkey feasts,\u201d where large quantities of food are spread out in gratitude to the monkeys for bringing good fortune.<\/p>\n<figure style=\"width: 308px\" class=\"wp-caption alignleft\"><img src=\"http:\/\/opentextbooks.concordia.ca\/wp-content\/uploads\/sites\/71\/2025\/07\/image51-1.jpg\" alt=\"Three macaques outside a temple in India.\" width=\"308\" height=\"261\" \/><figcaption class=\"wp-caption-text\">Figure 6.47: Because of important monkey-like figures in the Hindu religion, macaques are protected in India and often live near temples where they are fed by local peoples. Credit: <a href=\"https:\/\/commons.wikimedia.org\/wiki\/File:Macaque_India_4.jpg\">Macaque India 4<\/a> by <a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Mosmas\">Thomas Schoch<\/a> (<a href=\"https:\/\/commons.wikimedia.org\/wiki\/User:Mosmas\">Mosmas<\/a>) is under a <a href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/3.0\/\">CC BY-SA 3.0 License<\/a>.<\/figcaption><\/figure>\n<p class=\"import-Normal\">The people of Japan have coexisted with Japanese macaques for thousands of years, and so monkeys play key roles in both of the major Japanese religions. In the Shinto religion, macaques are thought of as messengers between the spirit world and humans, and monkey symbols are thought to be good luck. The other major religion in Japan is Buddhism, and monkeys play a role in symbolism of this religion as well. The \u201cThree Wise Monkeys\u201d who see no evil, speak no evil, and hear no evil derive from Buddhist iconography of monkeys.<\/p>\n<p class=\"import-Normal\">In Central and South America, monkeys feature often in Mayan and Aztec stories. In the Mayan creation story, the Popol Vuh, the \u201chero brothers,\u201d are actually a howler monkey and a spider monkey, who represent ancestors of humans in the story. In the Aztec religion, spider monkeys are associated with the god of arts, pleasure, and playfulness. A spider monkey is also represented in a Peruvian Nazca geoglyph, a large design made on the ground by moving rocks.<\/p>\n<p class=\"import-Normal\">In many of these regions today, the relationships between humans and nonhuman primates are complicated. The bushmeat and pet trades make these animals valuable at the expense of many animals\u2019 lives, and in some areas, nonhuman primates have become pests who raid crop fields and consume valuable foods. All of this has led to the development of a new subarea of anthropology called <strong><a class=\"glossary-term\" aria-haspopup=\"dialog\" aria-describedby=\"definition\" href=\"#term_1683_1440\">Ethnoprimatology<\/a><\/strong>, which involves studying the political, economic, symbolic, and practical relationships between humans and nonhuman primates.This field highlights the particular challenges for humans of having to coexist with animals with whom we share so much in common. It also provides insight into some of the challenges facing primate conservation efforts (see Appendix B: Primate Conservation).<\/p>\n<\/div>\n<h2 class=\"import-Normal\">Summary<\/h2>\n<p class=\"import-Normal\">The Order Primates is a diverse and fascinating group of animals united in sharing a suite of characteristics\u2014visual specialization, grasping hands and feet, large brains, and extended life histories\u2014that differentiates us from other groups of mammals. In this chapter, we surveyed the major taxonomic groups of primates, discussing where humans fit among our close relatives as well as discovering that primates are interesting animals in their own right. We discussed a range of key traits used to distinguish between the many taxa of living primates, including dietary, locomotor, and behavioural characteristics. Because of our long, shared evolutionary history with these animals, nonhuman primates provide a crucial resource for understanding our current biology. In Chapter 7, you will discover the fascinating and complex social behaviours of nonhuman primates, which provide further insight into our evolutionary biology.<\/p>\n<\/div>\n<h2>Expand your view<\/h2>\n<p>For a deeper look into primate conservation and contemporary issues of endangerment\/extinction see <a href=\"https:\/\/opentextbooks.concordia.ca\/explorations3\/back-matter\/appendix-b-primate-conservation\/\">Appendix B<\/a><\/p>\n<p>&nbsp;<\/p>\n<div class=\"learning-objectives\">\n<div class=\"textbox shaded\">\n<h2 class=\"import-Normal\">Review Questions<\/h2>\n<ul>\n<li class=\"import-Normal\">Why does the field of anthropology, a field dedicated to the study of humans, include the study of nonhuman animals? What can we learn from nonhuman primates?<\/li>\n<li class=\"import-Normal\">Why is it important to try to place taxa into a clade classification rather than groupings based on grade? Can you think of an example?<\/li>\n<li class=\"import-Normal\">One of the important goals of an introductory biological anthropology course is to teach you about your place in nature. What is the full taxonomic classification of humans, and what are some of the traits we have of each of these categories?<\/li>\n<li class=\"import-Normal\">When you have seen primates in person, did you observe any facial expressions, behaviours, or physical traits that seemed familiar to you? If so, which ones and why?<\/li>\n<li class=\"import-Normal\">Draw out a tree showing the major taxonomic group of primates described here, making sure to leave room in between each level. Underneath each taxon, list some of the key features of this group so that you can compare traits between groups.<\/li>\n<\/ul>\n<\/div>\n<h2 class=\"import-Normal\">Key Terms<\/h2>\n<p class=\"import-Normal\"><strong>Activity pattern<\/strong>: Refers to the time of day an animal is typically active.<\/p>\n<p class=\"import-Normal\"><strong>African clade<\/strong>: A grouping that includes gorillas, chimpanzees, bonobos, humans, and their extinct relatives.<\/p>\n<p class=\"import-Normal\"><strong>Analogy<\/strong>: When two or more taxa exhibit similar traits that have evolved independently, the similar traits evolve due to similar selective pressures. (Also sometimes called convergent evolution, parallel evolution, or homoplasy.)<\/p>\n<p class=\"import-Normal\"><strong>Ancestral trait<\/strong>: A trait that has been inherited from a distant ancestor.<\/p>\n<p class=\"import-Normal\"><strong>Arboreal<\/strong>: A descriptor for an organism that spends most of its time in trees.<\/p>\n<p class=\"import-Normal\"><strong>Asian clade<\/strong>: A grouping that includes orangutang and their extinct relatives.<\/p>\n<p class=\"import-Normal\"><strong>Bilophodont<\/strong>: Molar pattern of cercopithecoid monkeys in which there are usually four cusps that are arranged in a square pattern and connected by two ridges.<\/p>\n<p class=\"import-Normal\"><strong>Bipedalism<\/strong>: Walking on two legs.<\/p>\n<p class=\"import-Normal\"><strong>Brachiation<\/strong>: A form of locomotion in which the organism swings below branches using the forelimbs.<\/p>\n<p class=\"import-Normal\"><strong>Bunodont<\/strong>: Low, rounded cusps on the cheek teeth.<\/p>\n<p class=\"import-Normal\"><strong>Canines<\/strong>: In most primates, these are the longest of the teeth, often conical in shape and used as a weapon against predators or others of their species.<\/p>\n<p class=\"import-Normal\"><strong>Cathemeral<\/strong>: Active throughout the 24-hour period.<\/p>\n<p class=\"import-Normal\"><strong>Clade<\/strong>: A grouping based on ancestral relationships; a branch of the evolutionary tree.<\/p>\n<p class=\"import-Normal\"><strong>Cusps<\/strong>: The bumps on the chewing surface of the premolars and molars, which can be quite sharp in some species.<\/p>\n<p class=\"import-Normal\"><strong>Dental formula<\/strong>: The number of each type of tooth in one quadrant of the mouth, written as number of incisors: canines: premolars: molars.<\/p>\n<p class=\"import-Normal\"><strong>Derived trait<\/strong>: A trait that has been recently modified, most helpful when assigning taxonomic classification.<\/p>\n<p class=\"import-Normal\"><strong>Diastema<\/strong>: A space between the teeth, usually for large canines to fit when the mouth is closed.<\/p>\n<p class=\"import-Normal\"><strong>Dichromatic<\/strong>: Being able to see only blues and greens.<\/p>\n<p class=\"import-Normal\"><strong>Diurnal<\/strong>: Active during the day.<\/p>\n<p class=\"import-Normal\"><strong>Dry nose<\/strong>: The nose and upper lip are separated and the upper lip can move independently; sometimes referred to as a \u201chairy\u201d or \u201cmobile\u201d upper lip.<\/p>\n<p class=\"import-Normal\"><strong>Ethnoprimatology<\/strong>: A subarea of anthropology that studies the complexities of human-primate relationships in the modern environment.<\/p>\n<p class=\"import-Normal\"><strong>Evolutionary trade-off<\/strong>: When an organism, which is limited in the time and energy it can put into aspects of its biology and behaviour, is shaped by natural selection to invest in one adaptation at the expense of another.<\/p>\n<p class=\"import-Normal\"><strong>Faunivorous<\/strong>: Having a diet consisting entirely of animal matter: insects, eggs, lizards, etc.<\/p>\n<p class=\"import-Normal\"><strong>Folivore<\/strong>: Having a diet consisting primarily of leaves.<\/p>\n<p class=\"import-Normal\"><strong>Fovea<\/strong>: A depressed area in the retina at the back of the eye containing a concentration of cells that allow one to focus on objects very close to one\u2019s face.<\/p>\n<p class=\"import-Normal\"><strong>Frugivore<\/strong>: Having a diet consisting primarily of fruit.<\/p>\n<p class=\"import-Normal\"><strong>Generalized trait<\/strong>: A trait that is useful for a wide range of tasks.<\/p>\n<p class=\"import-Normal\"><strong>Grade<\/strong>: A grouping based on overall similarity in lifestyle, appearance, and behaviour.<\/p>\n<p class=\"import-Normal\"><strong>Grooming claw<\/strong>: A claw present on the second pedal digit in strepsirrhines.<\/p>\n<p class=\"import-Normal\"><strong>Gummivore<\/strong>: Having a diet consisting primarily of gums and saps.<\/p>\n<p class=\"import-Normal\"><strong>Heterodont<\/strong>: Having different types of teeth.<\/p>\n<p class=\"import-Normal\"><strong>Homology<\/strong>: When two or more taxa share characteristics because they inherited them from a common ancestor.<\/p>\n<p class=\"import-Normal\"><strong>Hone<\/strong>: When primates sharpen their canines by wearing them on adjacent teeth.<\/p>\n<p class=\"import-Normal\"><strong>Incisors<\/strong>: The spatula-shaped teeth at the front of the mouth.<\/p>\n<p class=\"import-Normal\"><strong>Insectivore<\/strong>: Having a diet consisting primarily of insects.<\/p>\n<p class=\"import-Normal\"><strong>Ischial callosities<\/strong>: Modified seat bones of the pelvis that are flattened and over which calluses form; function as seat pads for sitting and resting atop branches.<\/p>\n<p class=\"import-Normal\"><strong>Knuckle-walking<\/strong>: A form of quadrupedal movement used by <em>Gorilla<\/em> and <em>Pan<\/em> when on the ground, wherein the front limbs are supported on the knuckles of the hands.<\/p>\n<p class=\"import-Normal\"><strong>Life history<\/strong>: Refers to an organism\u2019s pace of growth, reproduction, lifespan, etc.<\/p>\n<p class=\"import-Normal\"><strong>Locomotion<\/strong>: How an organism moves around.<\/p>\n<p class=\"import-Normal\"><strong>Male bimaturism<\/strong>: Refers to the alternative reproductive strategies in orangutans in which males can delay maturation, sometimes indefinitely, until a fully mature, \u201cflanged\u201d male disappears.<\/p>\n<p class=\"import-Normal\"><strong>Molars<\/strong>: The largest teeth at the back of the mouth; used for chewing. In primates, these teeth usually have between three and five cusps.<\/p>\n<p class=\"import-Normal\"><strong>Monochromatic<\/strong>: Being able to see only in shades of light to dark, no colour.<\/p>\n<p class=\"import-Normal\"><strong>Monomorphic<\/strong>: When males and females of a species do not exhibit significant sexual dimorphism.<\/p>\n<p class=\"import-Normal\"><strong>Natal coat<\/strong>: Refers to the contrasting fur colour of baby leaf monkeys compared to adults.<\/p>\n<p class=\"import-Normal\"><strong>Nocturnal<\/strong>: Active at night.<\/p>\n<p class=\"import-Normal\"><strong>Olecranon process<\/strong>: Bony projection at the elbow end of the ulna.<\/p>\n<p class=\"import-Normal\"><strong>Opposable thumb <\/strong>or <strong>opposable big toe<\/strong>: Having thumbs and toes that go in a different direction from the rest of the fingers, allows for grasping with hands and feet.<\/p>\n<p class=\"import-Normal\"><strong>Pentadactyly<\/strong>: Having five digits or fingers and toes.<\/p>\n<p class=\"import-Normal\"><strong>Polymorphic colour vision<\/strong>: A system in which individuals of a species vary in their abilities to see colour. In primates, it refers to males being dichromatic and females being either trichromatic or dichromatic.<\/p>\n<p class=\"import-Normal\"><strong>Postorbital bar<\/strong>: A bony ring that surrounds the eye socket, open at the back.<\/p>\n<p class=\"import-Normal\"><strong>Postorbital closure\/plate<\/strong>: A bony plate that provides protection to the side and back of the eye.<\/p>\n<p class=\"import-Normal\"><strong>Prehensile tail<\/strong>: A tail that is able to hold the full body weight of an organism, which often has a tactile pad on the underside of the tip for improved grip.<\/p>\n<p class=\"import-Normal\"><strong>Premolars<\/strong>: Smaller than the molars, used for chewing. In primates, these teeth usually have one or two cusps.<\/p>\n<p class=\"import-Normal\"><strong>Quadrupedalism<\/strong>: Moving around on all fours.<\/p>\n<p class=\"import-Normal\"><strong>Rhinariums<\/strong>: Wet noses; resulting from naked skin of the nose which connects to the upper lip and smell-sensitive structures along the roof of the mouth.<\/p>\n<p class=\"import-Normal\"><strong>Sagittal crest<\/strong>: A bony ridge along the top\/middle of the skull, used for attachment of chewing muscles.<\/p>\n<p class=\"import-Normal\"><strong>Scent marking<\/strong>: The behaviour of rubbing scent glands or urine onto objects as a way of communicating with others.<\/p>\n<p class=\"import-Normal\"><strong>Semi-brachiation<\/strong>: A form of locomotion in which an organism swings below branches using a combination of forelimbs and prehensile tail.<\/p>\n<p class=\"import-Normal\"><strong>Sexually dimorphic<\/strong>: When a species exhibits sex differences in morphology, behaviour, hormones, and\/or colouration.<\/p>\n<p class=\"import-Normal\"><strong>Shearing crests<\/strong>: Sharpened ridges that connect cusps on a bilophodont molar.<\/p>\n<p class=\"import-Normal\"><strong>Specialized trait<\/strong>: A trait that has been modified for a specific purpose.<\/p>\n<p class=\"import-Normal\"><strong>Styloid process of ulna<\/strong>: A bony projection of the ulna at the end near the wrist.<\/p>\n<p class=\"import-Normal\"><strong>Tactile pads<\/strong>: Sensitive skin at the fingertips for sense of touch. Animals with a prehensile tail have a tactile pad on the underside of the tail as well.<\/p>\n<p class=\"import-Normal\"><strong>Tapetum lucidum<\/strong>: Reflecting layer at the back of the eye that magnifies light.<\/p>\n<p class=\"import-Normal\"><strong>Terrestrial<\/strong>: A descriptor for an organism that spends most of its time on the ground.<\/p>\n<p class=\"import-Normal\"><strong>Tetrachromatic<\/strong>: Having the ability to see reds, yellows, blues, greens, and ultraviolet.<\/p>\n<p class=\"import-Normal\"><strong>Tooth comb<\/strong> or <strong>dental comb<\/strong>: A trait of the front, lower teeth of strepsirrhines in which, typically, the four incisors and canines are long and thin and protrude outward.<\/p>\n<p class=\"import-Normal\"><strong>Trichromatic colour vision<\/strong>: Being able to distinguish yellows and reds in addition to blues and greens.<\/p>\n<p class=\"import-Normal\"><strong>Vertical clinging and leaping<\/strong>: A locomotor pattern in which animals are oriented upright while clinging to vertical branches, push off with hind legs, and land oriented upright on another vertical branch.<\/p>\n<p class=\"import-Normal\"><strong>Y-5 molar<\/strong>: Molar cusp pattern in which five molar cusps are separated by a \u201cY\u201d-shaped groove pattern.<\/p>\n<h2 class=\"import-Normal\">For Further Exploration<\/h2>\n<p class=\"import-Normal\"><a href=\"https:\/\/animaldiversity.org\/accounts\/Primates\/specimens\/\">Animal Diversity Web<\/a>. This website is hosted by the Zoology Department at the University of Michigan. It has photographs of skulls, teeth, hands, arms, and feet of many primate species.<\/p>\n<p class=\"import-Normal\"><a href=\"https:\/\/www.eskeletons.org\">eSkeletons<\/a>. This website is hosted by the Department of Anthropology at University of Texas, Austin. It is an interactive website where you can compare specific bones from different species of primates.<\/p>\n<p class=\"import-Normal\">Fleagle, John G. 2013. <em>Primate Adaptation and Evolution<\/em>. Third edition. San Diego: Academic Press.<\/p>\n<p class=\"import-Normal\">Fuentes, Agust\u00edn, and Kimberley J. Hockings. 2010. \u201cThe Ethnoprimatological Approach in Primatology.\u201d <em>American Journal of Primatology<\/em> 72 (10): 841\u2013847.<\/p>\n<p class=\"import-Normal\">Rowe, Noel. 1996. <em>Pictorial Guide to the Living Primates<\/em>. Charlestown, RI: Pogonias Press.<\/p>\n<p class=\"import-Normal\">Whitehead, Paul F., William K. Sacco, and Susan B. Hochgraf. 2005. <em>A Photographic Atlas for Physical Anthropology<\/em>. Englewood, CO: Morton Publishing.<\/p>\n<h2>References<\/h2>\n<p class=\"import-Normal\">Balolia, Katharine L., Christophe Soligo, and Bernard Wood. 2017. \u201cSagittal Crest Formation in Great Apes and Gibbons.\u201d <em>Journal of Anatomy<\/em> 230 (6): 820\u2013832.<\/p>\n<p class=\"import-Normal\">Bininda-Emonds, Olaf R., Marcel Cardillo, Kate E. Jones, Ross D. E. MacPhee, Robin M. D. Beck, Richard Grenyer, Samantha A. Price, Rutger A. Vos, John L. Gittleman, and Andy Purvis. 2007. \u201cThe Delayed Rise of Present-Day Mammals.\u201d <em>Nature<\/em> 446 (7135): 507\u2013512.<\/p>\n<p class=\"import-Normal\">Burton, Michael L., Carmella C. Moore, John W. M. Whiting, A. Kimball Romney, David F. Aberle, Juan A. Barcelo, Malcolm M. Dow, et al. 1996. \u201cRegions Based on Social Structure.\u201d <em>Current Anthropology<\/em> 37 (1): 87\u2013123.<\/p>\n<p class=\"import-Normal\">Chivers, David J., and C. M. Hladik. 1980. \u201cMorphology of the Gastrointestinal Tract in Primates: Comparisons with Other Mammals in Relation to Diet.\u201d <em>Journal of Morphology<\/em> 166 (3): 337\u2013386.<\/p>\n<p class=\"import-Normal\">Clutton-Brock, T. H., and Paul H. Harvey. 1980. \u201cPrimates, Brains, and Ecology.\u201d <em>Journal of Zoology<\/em> 190 (3): 309\u2013323.<\/p>\n<p class=\"import-Normal\">Dunbar, Robin I. M. 1998. \u201cThe Social Brain Hypothesis.\u201d <em>Evolutionary Anthropology<\/em> 6 (5): 178\u2013190.<\/p>\n<p class=\"import-Normal\">Ebersberger, Ingo, Dirk Metzler, Carsten Schwarz, and Svante P\u00e4\u00e4bo. 2002. \u201cGenomewide Comparison of DNA Sequences Between Humans and Chimpanzees.\u201d <em>American Journal of Human Genetics<\/em> 70 (6): 1490\u20131497.<\/p>\n<p class=\"import-Normal\">Jameson, Natalie M., Zhuo-Cheng Hou, Kirstin N. Sterner, Amy Weckle, Morris Goodman, Michael E. Steiper, and Derek E. Wildman. 2011. \u201cGenomic Data Reject the Hypothesis of a Prosimian Primate Clade.\u201d <em>Journal of Human Evolution<\/em> 61 (3): 295\u2013305.<\/p>\n<p class=\"import-Normal\">Kawamura, Shoji, Chihiro Hiramatsu, Amanda D. Melin, Colleen M. Schaffner, Filippo Aureli, and Linda M. Fedigan. 2012. \u201cPolymorphic Color Vision in Primates: Evolutionary Considerations.\u201d In <em>Post-Genome Biology of Primates<\/em>, edited by H. Irai, H. Imai, and Y. Go, 93\u2013120. Tokyo: Springer.<\/p>\n<p class=\"import-Normal\">Matsui, Atsushi, Felix Rakotondraparany, Isao Munechika, Masami Hasegawa, and Satoshi Horai. 2009. \u201cMolecular Phylogeny and Evolution of Prosimians Based on Complete Sequences of Mitochondrial DNAs.\u201d <em>Gene<\/em> 441 (1\u20132): 53\u201366.<\/p>\n<p class=\"import-Normal\">Nater, Alexander, Maja P. Mattle-Greminger, Anton Nurcahyo, Matthew G. Nowak, Marc de Manuel, Tariq Desai, Colin Groves, et al. 2017. \u201cMorphometric, Behavioral, and Genomic Evidence for a New Orangutan Species.\u201d <em>Current Biology<\/em> 27 (22): 3487\u20133498.<\/p>\n<p class=\"import-Normal\">Pozzi, Luca, Jason A. Hodgson, Andrew S. Burrell, Kirstin N. Sterner, Ryan L. Raaum, and Todd R. Disotell. 2014. \u201cPrimate Phylogenetic Relationships and Divergence Dates Inferred from Complete Mitochondrial Genomes.\u201d <em>Molecular Phylogenetics and Evolution<\/em> 75: 165\u2013183.<\/p>\n<p class=\"import-Normal\">Remis, Melissa J. 1997. \u201cWestern Lowland Gorillas (<em>Gorilla gorilla gorilla<\/em>) as Seasonal Frugivores: Use of Variable Resources.\u201d <em>American Journal of Primatology<\/em> 43 (2): 87\u2013109.<\/p>\n<p class=\"import-Normal\">Richmond, Brian G., David R. Begun, and David S. Strait. 2001. \u201cOrigin of Human Bipedalism: The Knuckle\u2010Walking Hypothesis Revisited.\u201d <em>American<\/em> <em>Journal of Physical Anthropology<\/em> 116 (S33): 70\u2013105.<\/p>\n<p class=\"import-Normal\">Robson, Shannen L., Carel P. van Schaik, and Kristen Hawkes. 2006. \u201cThe Derived Features of Human Life History.\u201d In <em>The Evolution of Human Life History, edited by Kristen Hawkes and Richard R. Paine, <\/em>17\u201344. Santa Fe: SAR Press.<\/p>\n<p class=\"import-Normal\">Scally, Aylwyn, Julien Y. Dutheil, LaDeana W. Hillier, Gregory E. Jordan, Ian Goodhead, Javier Herrero, Asger Hobolth, et al. 2012. \u201cInsights into Hominid Evolution from the Gorilla Genome Sequence.\u201d <em>Nature<\/em> 483 (7388): 169\u2013175.<\/p>\n<p class=\"import-Normal\">Schneider, Horacio, and Iracilda Sampaio. 2015. \u201cThe Systematics and Evolution of New World Primates: A Review.\u201d <em>Molecular Phylogenetics and Evolution<\/em> 82 (B): 348\u2013357.<\/p>\n<p class=\"import-Normal\">Setchell, Joanna M., Phyllis C. Lee, E. Jean Wickings, and Alan F. Dixson. 2001. \u201cGrowth and Ontogeny of Sexual Size Dimorphism in the Mandrill (<em>Mandrillus sphinx<\/em>).\u201d <em>American Journal of Physical Anthropology<\/em> 115 (4): 349\u2013360.<\/p>\n<p class=\"import-Normal\">Utami, Sri Suci, Beno\u00eet Goossens, Michael W. Bruford, Jan R. de Ruiter, and Jan A. R. A. M. van Hooff. 2002. \u201cMale Bimaturism and Reproductive Success in Sumatran Orang-utans.\u201d <em>Behavioral Ecology<\/em> 13 (5): 643\u2013652.<\/p>\n<p class=\"import-Normal\">Vasey, Natalie. 2006. \u201cImpact of Seasonality and Reproduction on Social Structure, Ranging Patterns, and Fission\u2013Fusion Social Organization in Red Ruffed Lemurs.\u201d In <em>Lemurs: Ecology and Adaptation<\/em>, edited by Lisa Gould and Michelle L. Sauther, 275\u2013304. New York: Springer.<\/p>\n<p class=\"import-Normal\">Wright, Patricia C. 1999. \u201cLemur Traits and Madagascar Ecology: Coping with an Island Environment.\u201d <em>American Journal of Physical Anthropology<\/em> 110 (S29): 31\u201372.<\/p>\n<h2>Acknowledgements<\/h2>\n<p class=\"import-Normal\">The author would very much like to thank the editors for the opportunity to contribute to this textbook, along with anonymous reviewers who provided useful feedback on earlier drafts of this chapter. She would particularly like to thank Karin Enstam Jaffe for her support and encouragement during the writing of this chapter and its revision. Most of all, the author would like to thank all of the Introduction to Biological Anthropology students that she has had over the years who have listened to her lecture endlessly on these animals that she finds so fascinating and who have helped her to hone her pedagogy in a field that she loves.<\/p>\n<\/div>\n<\/div>\n<\/div><button><span aria-hidden=\"true\">&times;<\/span><span class=\"screen-reader-text\">Close definition<\/span><\/button><\/div><\/template><\/div>","protected":false},"author":96,"menu_order":18,"template":"","meta":{"pb_show_title":"on","pb_short_title":"","pb_subtitle":"","pb_authors":["jonathan-m-g-perry","stephanie-l-canington"],"pb_section_license":""},"chapter-type":[],"contributor":[93,94],"license":[],"class_list":["post-1683","chapter","type-chapter","status-publish","hentry","contributor-jonathan-m-g-perry","contributor-stephanie-l-canington"],"part":20,"_links":{"self":[{"href":"https:\/\/opentextbooks.concordia.ca\/explorations3\/wp-json\/pressbooks\/v2\/chapters\/1683","targetHints":{"allow":["GET"]}}],"collection":[{"href":"https:\/\/opentextbooks.concordia.ca\/explorations3\/wp-json\/pressbooks\/v2\/chapters"}],"about":[{"href":"https:\/\/opentextbooks.concordia.ca\/explorations3\/wp-json\/wp\/v2\/types\/chapter"}],"author":[{"embeddable":true,"href":"https:\/\/opentextbooks.concordia.ca\/explorations3\/wp-json\/wp\/v2\/users\/96"}],"version-history":[{"count":6,"href":"https:\/\/opentextbooks.concordia.ca\/explorations3\/wp-json\/pressbooks\/v2\/chapters\/1683\/revisions"}],"predecessor-version":[{"id":1783,"href":"https:\/\/opentextbooks.concordia.ca\/explorations3\/wp-json\/pressbooks\/v2\/chapters\/1683\/revisions\/1783"}],"part":[{"href":"https:\/\/opentextbooks.concordia.ca\/explorations3\/wp-json\/pressbooks\/v2\/parts\/20"}],"metadata":[{"href":"https:\/\/opentextbooks.concordia.ca\/explorations3\/wp-json\/pressbooks\/v2\/chapters\/1683\/metadata\/"}],"wp:attachment":[{"href":"https:\/\/opentextbooks.concordia.ca\/explorations3\/wp-json\/wp\/v2\/media?parent=1683"}],"wp:term":[{"taxonomy":"chapter-type","embeddable":true,"href":"https:\/\/opentextbooks.concordia.ca\/explorations3\/wp-json\/pressbooks\/v2\/chapter-type?post=1683"},{"taxonomy":"contributor","embeddable":true,"href":"https:\/\/opentextbooks.concordia.ca\/explorations3\/wp-json\/wp\/v2\/contributor?post=1683"},{"taxonomy":"license","embeddable":true,"href":"https:\/\/opentextbooks.concordia.ca\/explorations3\/wp-json\/wp\/v2\/license?post=1683"}],"curies":[{"name":"wp","href":"https:\/\/api.w.org\/{rel}","templated":true}]}}